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Records of the Museums and Art Galleries
of the Northern Territory

Volume 24

December 2008

The Beagle, Records of the Museums and Art Galleries of the Northern Territory

(formerly ‘Records of the Northern Territory Museum of Arts and Sciences )

EDITORIAL COMMITTEE

R. C. Willan Editor D. Carment

C. J. Glasby Associate Editor H. K. Larson

D. Megirian
P. Murray
P. Short

The Beagle is a refereed journal published by the Museums and Art Galleries of the Northern Territory to disseminate the results
of research in the following areas:

Systematic and other studies of the terrestrial, marine and freshwater flora and fauna of the Northern Territory, tropical
Australia, Southeast Asia and the Indo-Pacific;

Australian Aboriginal, Southeast Asian and Oceanic art, material culture and archaeology;

Northern Territory and Oceanic history and archaeology.

The Beagle is published once or twice a year, depending upon the material available.

All contributions to The Beagle are reviewed by two referees and, where possible, at least one is internationally-based.

Whilst articles for The Beagle will normally be 10-50 pages in length, shorter communications, notes and review articles may
also be acceptable. Longer articles, significant works or substantial revisions, which form integral studies, may be considered for
separate publication as a Supplement. Prospective authors should contact the Editor, Academic Publications.

The Beagle may be obtained by subscription or by exchange. The subscription rate for one year for individuals and institutions
is S66.00 (includes postage and GST). All orders, subscriptions, back numbers and exchange enquiries should be addressed to the
Library Manager, Museum and Art Gallery Northern Territory, GPO Box 4646, Darwin NT 0801, AUSTRALIA, or e-mail library.
dam@nt.gov.au

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Twenty-five offprints are provided free for each published paper. Additional offprints may be ordered when returning proofs.

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GPO Box 4646, Darwin NT0801, AUSTRALIA
E-mail: richard. willan@nt.gov. an

ISSN 1833-7511

©Museums and Art Galleries of the Northern Territory, 2008.
Printed by the Government Printing Office of the Northern Territory

DARWIN200

MUSEUM
AND ART
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NORTHERN

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Front cover: Pencil-blue butterflies of the Candalides absimilis species group from eastern Australia (see Braby, pages 33-54).
Lower left: C. absimilis absimilis; all others: C. absimilis edwardsi. Photographs by Michael Braby.

The Beagle

Records of the Museums and Art Galleries
of the Northern Territory

Volume 24, December 2008
CONTENTS

BERRA, T. M. - Charles Darwin’s paradigm shift.1

DURETTO, M. F. — A reassessment of Boronia (Rutaceae) in the Northern Territory with a key to species,

the description of one new species and the reduction, in synonymy, of another species.7

PAXTON, H and SAFARIK, M. - Jaw growth and replacement in Diopatra aciculata (Annelida: Onuphidae).15

SALGADO KENT, C. P. and McGUINNESS, K. A. - Feeding selectivity of sesarmid crabs from

northern Australian mangrove forests.23

BRABY, M. F. - Taxonomic review of Candalides absimilis (C. Felder, 1862) and C. margarita (Semper, 1879)
(Lepidoptera: Lycaenidae), with descriptions of two new subspecies.33

BAEHR, M. - Two new species of the genus Pseudaptinus Castelnau from northern Australia

(Insecta: Coleoptera: Carabidae: Zuphiinae). 55

KOTT, P. - Biogeographic implications of Ascidiacea (Tunicata) from the Wessel Islands (Arafura Sea).63

THORBURN, D. C. and ROWLAND, A. J. - Juvenile bull sharks Carcharhinus leucas (Valenciennes, 1839)

in northern Australian rivers.79

M0LLER, P. R. and SCHWARZHANS, W. - Review of the Dinematichthyni (Teleostci: Bythitidae)

of the Indo-west Pacific. Part IV. Dinematichthys and two new genera with descriptions of nine new species.87

LARSON, H. K. - A new species of the gudgeon Bostrychus (Teleostei: Gobioidei: Eleotridae), from

peninsular Malaysia. 147

MOHD-AZLAN, J., TAHA, S. H., LAMAN, C. J. M. and ABDULLAH, M. T. - Diversity of bats at

two contrasting elevations in a protected dipterocarp forest in Sarawak, Borneo.151

Short communication

TENNENT, W. J. - A previously unpublished record of Liphyra brassolis Westwood, 1864

(Lepidoptera: Lycaenidae: Miletinae) from Vella Lavella, New Georgia group, Solomon Islands.157

Corrigendum

HORNER, P. - Corrigendum to Horner, P. and Adams, M. (2007). A molecular systematic assessment of
species boundaries in Australian Cryptoblepharus (Reptilia: Squamata: Scincidae) - a case study for the
combined use of allozymes and morphology to explore cryptic biodiversity. The Beagle , Records of the
Museums and Art Galleries of the Northern Territory, Supplement 3: 1-19,

159

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24 : 1-5

Charles Darwin’s paradigm shift

TIM M. BERRA

Research Associate, Museum and Art Gallery Northern Territory

and

Department of Evolution, Ecology> and Organismal Biology,

The Ohio State University, Mansfield, Ohio 44906, USA
berra. l@osu.edu

ABSTRACT

The publication ofDarwin’s On the Origin oj Species in 1859 created a paradigm shift from creation to evolution. Darwin
showed that humans are part ot nature, not above it. and that all animal life, including human, is related by descent
from a common ancestor. His mechanism of evolution via natural selection is a powerful creative force that provided
an explanation for the diversity of life. This dramatic change in world view from supematuralism to methodological
naturalism has allowed staggering scientific advances in the past 150 years that transcend science and impact on the
human psyche.

Keywords: Charles Darwin, evolution, natural selection, naturalism, paradigm shift.

INTRODUCTION

Charles Darwin (1809-1882) was an extraordinary man
by any standard. The theory of evolution by natural selection
as elaborated in his book On the origin of species (1859) is
considered by historians and philosophers of science to be
one of the most important ideas ever had by the human mind
(Dennett 1995). Before exploring this grandiose statement,
a brief review ofDarwin’s life and scientific accomplish¬
ments is in order. Then I will address the implications of
his very useful insight that extended beyond science and
profoundly impacted on the human mind.

An outline of Darwin’s Life

Charles Darwin was bom into a wealthy English family
on 12 February 1809. His father, Robert Waring Darwin
(1766-1848), was a prominent physician as was his grand¬
father Erasmus Darwin (1731-1802). His mother was
Susannah Wedgwood (1764-1817), the daughter of Josiah
Wedgwood (1730-1795), the pottery manufacturer and
entrepreneur, who was a close friend of Erasmus Darwin.

Darwin’s father sent Charles to medical school at
Edinburgh University in 1825 and removed him in 1827
when it became obvious that Charles was not interested in
a medical career. Robert Darwin then decided that Charles
should study to be a clergyman in the Church of England,
and sent him to Cambridge University in 1828. Charles
graduated 10"' in his class in 1831 and then received an
invitation orchestrated by his professor, John Stevens Hens-
low (1796-1861), to be an unpaid naturalist-companion to
Captain Robert FitzRoy (1805-1865) on a surveying voyage
around the world on I I.M.S. Beagle (1831-1836). Darwin
later described this opportunity as “the first real training or
education of my mind”.

Upon return from the nearly five-year Beagle voyage,
Darwin found that he was accepted as a serious scientist,
and he had no desire to become a clergyman. He began
working on the specimens collected on the voyage. He
married his first cousin, Emma Wedgwood (1808-1896),
and they eventually moved from London to Down House
in Kent (Fig. 1). They had 10 children, seven of whom sur¬
vived to adulthood. After the voyage, he was often ill, but
nevertheless, highly productive. He entered his ideas about
how species form in a series of notebooks This included a

Fig. 1. At rear of montage, wedding portrait of Charles Darwin
(watercolour) in 1840 at age 30 by George Richmond. At centre,
1IMS Beagle drawing by American artist Samuel L. Margolies
(1897-1974) from Dibncr (1960) in the Bumdy Collection at the
Huntington Library, San Marino, CA. Used with permission of the
Huntington Library. In foreground, one of the last photographs of
Charles Darwin, by Elliot and Fiy.

T. M. Berra

branching, tree-like diagram that reflected the common ori¬
gin and relatedness of organisms. This first evolutionary tree
showed that classification should be genealogical. However,
he kept his revolutionary ideas private for 20 years except
for his closest scientific colleagues: geologist Charles Lyell
(1797-1875), botanist Joseph Dalton Hooker (1817-1911),
zoologist Thomas Henry Huxley (1825-1895), and his
American botanist correspondent at Harvard University,
Asa Gray (1810-1888). In 1858 Darwin received of a let¬
ter from naturalist Alfred Russel Wallace (1823-1913),
who, like Darwin, was inspired by the writings of Thomas
Malthus (1766-1834). Wallace outlined ideas nearly iden¬
tical to Darwin’s. This letter and urging from Lyell and
Hooker prompted him to complete and publish On the
Origin of Species in 1859. Darwin continued to do experi¬
ments and publish on a variety of topics right up to the time
of his death. He died of heart disease on 19 April 1882 and
was laid to rest with pomp and ceremony in Westminster
Abby a few feet from Isaac Newton. Further details can be
pursued in three of the most comprehensive biographies of
Darwin (Desmond and Moore 1991; Browne 1995, 2002),
a concise biography (Berra 2009), and, of course, Darwin’s
autobiography (Barlow 1958).

SYNOPSIS OF DARWIN’S SCIENTIFIC
ACHIEVEMENTS

The educated citizen is generally aware of The origin
and Darwin’s account of his voyage around the world in
H.M.S. Beagle through his book now universally known
as The voyage of the Beagle. These two books have never
been out of print.

Most people are surprised to learn that Darwin also made
many other major contributions to geology, zoology, and
botany through his observations, experiments and writings.
His books have been chronicled (Berra 2009), so I will
just briefly outline the breadth of his influence. Darwin
explained how coral reefs form (1842) and contributed to
geological observations on earth movements (1844) and
deformation theory of metamorphic rock (1846). In a pio¬
neering four-volume work that took eight years to complete,
he described all known barnacle species, fossil and living
(1851-1854). Darwin explained how orchids are fertilized
by insects (1862), how plants climb (1865), and catalogued
the bewildering amount of variation in domestic plants and
animals (1868). He explained human origins and sexual
selection in ways never before articulated (1870-71), and
discussed human and animal emotions in similar terms
(1872). The latter work was one of the first books to use
photographs to illustrate a point.

Darwin showed how insectivorous plants on impov¬
erished soils utilise nitrogen-rich insects (1875), and
demonstrated that the offspring of cross-fertilised plants
were more numerous and vigorous than self-fertilised ones
(1876, 1877). His observations of climbing plants laid the

foundation for the field of plant growth hormones (1880),
and his work on earthworms (1881) is a classic study in
ecology. Any one of these achievements could constitute a
life’s work for most scientists.

DARWIN’S LEGACY

Darwin was bom and educated at a time when special
creation was the prevailing scientific view. That is, God cre¬
ated the universe and all species a few thousand years ago,
and they were unchangeable. “Revelation”, not research,
provided this view. Darwin began the Beagle voyage with
this belief. During his lifetime the age of the earth was in¬
creasingly recognised as ancient as suggested by Georges
Cuvier (1769-1832) and Charles Lyell (Bowler 1984;
Larson 2004). Observations made during the voyage made
him question the Genesis creation myth and immutability of
species. He found marine fossils thousands of feet above sea
level and reasoned that the land had been elevated by earth
movements, not inundated in a great biblical flood. The
fossil mammals he uncovered in South America resembled
living mammals from the same area. He wondered why this
should be if each species were specially created. Extinction
was hardly recognised in those days. Why did the animals
on islands off continental areas resemble those of the nearest
land mass if each species were created in place? Why were
there so many species in an island group that looked very
similar but with slight differences from island to island?
It is as if “one species had been taken and modified for
different ends”, he wrote in Voyage of the Beagle. None of
these things made sense from a creationist perspective. In
1844 he wrote to Hooker that “I am almost convinced (quite
contrary to the opinion I started with) that species are not
(it is like confessing a murder) immutable.”

The elegant simplicity of Darwin’s reasoning can be
distilled as follows. There is variation in nature, many more
offspring are generated than can survive, therefore there is a
struggle for life in which favorable variations arc preserved
and unfavorable variations arc removed. This leads to evolu¬
tion which he defined as “descent with modification” and to
the formation of new species. Nature is doing the selecting
for the forms best adapted to a particular environment so he
called the process natural selection as opposed to artificial
selection that breeders impose. We now know that muta¬
tion, chromosomal rearrangements, sexual reproduction,
etc. are the sources of genetic variation, but Darwin had no
knowledge of such topics. Today we can speak of “descent
with modification” as “a change in gene frequency”, and
natural selection is simply “differential reproduction”, that
is, one genetic variant leaves more offspring than another
(Berra 1990). Darwin borrowed the expression “survival of
the fittest” from economist Herbert Spencer (1820-1903).
Evolutionary fitness means reproductive fitness. In modem
terms, the fittest is the one who gets the most genes into

Charles Darwin’ paradigm shift

Fig. 2. Cartoon of Charles Darwin as a monkey, from Fun,
16 November 1872, just after The expression of the emotions in man
and animals was published. Many other similar personal attacks were
published during his lifetime.

the next generation, not necessarily the biggest or strongest
individual.

By the time of Darwin's death in 1882, most scientists of
the world had accepted the concept of common descent, but
some were still skeptical of natural selection as a creative
mechanism (Bowler 1984). The public was less accepting
(Fig. 2).

The publication of On the origin of species on 24 No¬
vember 1859 precipitated one of those rare events in the
history of science, a paradigm shift. Philosopher Thomas
Kuhn used this term to refer to the replacement of one world
view by another (Kuhn 1962). Examples ofa paradigm shift
in science include the replacement of the earth-centered
Ptolemaic system by the sun-centered Copemican system
and the replacement of Newtonian physics by relativity
and quantum physics.

Darwin’s work neatly dove-tailed into the wider pattern
of scientific advances that were occurring during his life.
Lyell and others had provided the necessary' geological time
for evolution to operate. The writings of Malthus, Spencer,
Wallace, and many others help set the evolutionary stage.
By 1859 evolution by natural selection was an idea whose
time had come. Darwin and the publication of The origin
closed the deal. Darwin changed the way humans view their
place in nature. He showed that humans were not above
nature, but pail of it. He supplied the explanation for the
great diversity of life and showed that all life, including

human, is related by descent from a common ancestor. His
explanation of evolution via natural selection is the basis
of all of biology and its applied subdisciplines of medicine,
agriculture, and biotechnology. No other biologist in the
history of our species has had an impact of this magni¬
tude. In the words of the eminent geneticist Theodosius
Dobzhansky, “Nothing in biology makes sense except in
the light of evolution” (Dobzhansky 1973).

The paradigm shift from creation to evolution has moved
intellectual endeavors from untestable belief to rational un¬
derstanding that flows from the scientific method. This, in
turn, has allowed a vast array of advances in knowledge.

DARWINIAN IMPLICATIONS

One of the attributes of a powerful scientific theory is
that it enables future research and understanding. Darwin¬
ian or evolutionary medicine as formulated by Nesse and
Williams (1996) explains how some disease symptoms,
such as fever, may be a response favored by natural selec¬
tion as a defense against pathogens. Some genetic diseases
such as sickle cell anaemia may allow differential survival
of its victims in malarial zones, a phenomenon called a
balanced polymorphism (Berra 1990). Evolutionary think¬
ing explains the arms race waged by pathogens and hosts
that prevents either from being completely eliminated.
The development of antibiotic resistant bacteria through
the flagrant overuse of antibiotics is easily explained by
Darwinian reasoning. A drug kills the susceptible bacteria
leaving bacteria with a pre-existing resistant mutation to
build up the next generation. Then when you actually need
the antibiotic for a bacterial infection, you find that the drug
is ineffective. This is evolution, pure and simple.

A similar process occurs in agriculture with the over
application of pesticides and the formation of pesticide
resistant pathogens, insects, and noxious plants. Austral¬
ians are very familiar with the myxomatosis versus rabbit
“anus race” whereby the virus initially killed 99 percent
of the rabbits, but given enough time the surviving rabbits
returned in force as the virus evolved in the direction of less
virulence and the rabbits were selected for more resistance
to the virus (Berra 1998).

Evolutionary psychology and evolutionary ethics, as
explored by Barkow et al. (1992) and popularised by Wright
(1994) help explain the origin of morality. Peacemaking
among non-human primates by the calming effect of mutual
grooming to diffuse aggression may be seen as the precur¬
sor of what became morality in humans (de Waal 1989).
Modem religions are recent human inventions - a mere
few thousand years old. The antecedents of morality, on the
other hand, clearly evolved before humanity as reflected in
the empathy exhibited by bonobos (Pan paniscus ) and the
reciprocity of chimpanzees (P. troglodytes) (de Waal 2005).
Kin selection, whereby an individual sacrifices for a close
genetic relative, makes sense in an evolutionary context

T. M. Berra

because some of the same genes of the individual making
the sacrifice will be passed on by the kin who survives.
This is referred to as inclusive fitness by Hamilton (1972).
Realisation that humans share kinship with all animal life
has helped to raise consciousness of how we treat other
animals (Singer 1977).

The ancestry of the AIDS virus, HIV-1 (human immu¬
nodeficiency virus-1) has been traced to SIVcpz (simian
immunodeficiency virus) carried by our closest living rela¬
tive, the chimpanzees. Pan troglodytes (Bailes et al. 2003).
This is not surprising from an evolutionary perspective.
Somewhere in high school today there is a student who
may contribute to the control of the AIDS epidemic. What
chance of that would there be if creationism were taught
as science in high school?

Even religion is now being explained as having an
evolutionary origin as a natural phenomenon once the brain
evolved a critical mass and complexity (Dennett 2006).
Bloch (2008) suggested that the evolution of imagination
was a requisite for the emergence of religion which he con¬
sidered a logical extension of human sociality. This occurred
about the time of the Upper Palaeolithic ''revolution" as
manifested by an explosion of image-making and cultural
transformations (White 2003). Acceptance of authority
necessary for group cohesion and survival enforced by tool
use and language combined with confusion of cause and
effect and coincidences can result in the establishment of a
belief that becomes dominate in a culture (Wolpcrt 2007).

Those whose religion requires a literal interpretation of
the Bible fear that the paradigm shift from supernatural ism
to methodological naturalism threatens their beliefs. The
1925 Scopes trial, nicknamed "monkey trial” and “trial of
the century'” in Dayton, Tennessee, has come to symbolise
the struggle of religion against science in popular culture
that later inspired the play and film Inherit the wind (Larson
1977). Such creationists arc particularly vocal in America
which has a long standing tradition of anti-intellectual ism
(Numbers 1992; Pigliucci 2002). This has resulted in a
series of creationist legal challenges to evolution which
have been decided in favor of evolution (Berra 1990). The
most recent of these was the Intelligent Design creationist
challenge of the Dover, Pennsylvania, School Board. The
Intelligent Design creationist philosophy that life is too
complex to have arisen by natural means and therefore had
a supernatural origin has been critiqued in Pennock (2001)
and exposed as a threat to science education by Forrest
and Gross (2004). In the conclusion of his decision Judge
John E. Jones III determined that the school board’s policy
of teaching Intelligent Design violated the Establishment
Clause of the First Amendment [separation of church and
state] of the U.S. Constitution, and he wrote, “...in making
this determination, we have addressed the seminal question
whether Intelligent Design is science (Jones 2005). We
have concluded that it is not, and moreover that Intelligent
Design cannot uncouple itself from its creationist, and thus
religious, antecedents.” He further wrote, "The breathtaking

inanity of the board’s decision is evident when considered
against the factual backdrop which has now been fully
revealed through this trial.” Padian (2007) reviewed three
books based on the Dover trial.

Biotechnology, whether in the form of genetically
modified crops, designer drugs, gene therapy, or the hu¬
man genome project all derive from Darwin’s profound
insight. Darwin had no knowledge of genes, chromosomes,
or how inheritance worked. This required additional input
from the understanding of Gregor Mendel’s (1822-1884)
genetic work.

The modem evolutionary synthesis grew from Darwin’s
explanation of natural selection and Mendel’s demonstration
of inheritance augmented by the research of mathematically
oriented population geneticists such as J.B.S. Haldane,
Ronald Fisher, Sewall Wright, Thomas Hunt Morgan,
Theodosius Dobzhansky, palaeontologist George Gaylord
Simpson, botanist G. Ledyard Stebbins, Jr., biologist Julian
Huxley (T.H. Huxley’s grandson), and the most important
evolutionary biologist since Darwin, Ernst Mayr. This fu¬
sion of knowledge moved evolutionary science forward
to the middle of the 20th century (Larson 2004). James D.
Watson’s and Francis Crick’s 1953 demonstration that the
molecular structure of DNA allowed for genetic coding
was a huge breakthrough that ultimately made it possible
to sequence the three billion chemical base pairs that com¬
pose the human genome and identify the approximately
20,000-25,000 genes in human DNA (Lander et al. 2001;
Venter et al. 2001).

Recent discoveries in evolutionary developmental biol¬
ogy, known as evo-devo, have shown that very similar genes
are present in very dissimilar animals. These body-shaping
genes are controlled by DNA switches called enhancers that
turn them on or off at various times in development. Such
enhancers are a major factor in the evolution of anatomy
(Carroll 2005). The above examples are just a sample of
the benefits to society that flow directly from the creative
power of Charles Darwin’s theory of evolution by means
of natural selection.

The paradigm shift instigated by Darwin has made
obvious the superiority of the scientific method as a means
of understanding the world around us. It is ironic that the
legacy of a man once destined for the church has been to
replace supernaturalism with methodological naturalism.

ACKNOWLEDGMENTS

I am grateful to the Director of the Museums and Art
Galleries of the Northern Territory, Anna Malgorezwicz,
and to the Editor of the Beagle , Richard Willan, for the
invitation to write this article in celebration of the bicenten¬
nial of Charles Darwin’s birth and the 150th anniversary
of the publication of On the origin of species in 2009. This
manuscript benefited greatly by a critique from Edward J.
Larson.

Charles Darwin’ paradigm shift

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Accepted 1 May 2008

■

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24 : 7-13

A reassessment of Boronia (Rutaceae) in the Northern Territory with a key to
species, the description of one new species and the reduction, in synonymy, of

another species

Marco F. Duretto

Tasmanian Herbarium, Tasmanian Museum and Art Gallery, Private Bag 4, Hobart, Tasmania 7001, AUSTRALIA.

Email: marco.duretto@tmag.tas.gov.au

ABSTRACT

Boronia (Rutaceae) is reassessed for the Northern Territory of Australia. Three informal names, B. aff. laxa 1 (Northern
Plateau, Arnhem Land), B. aff. laxa 2 (Nabarlek, Arnhem Land) and B. aff. prolixa 1 (Red Lily Lagoon, Arnhem Land),
are re-evaluated in the light of the significant number of new collections made over the previous decade. Ol the three,
only one, B. aff. laxa 2 (Nabarlek, Arnhem Land), is considered to be worthy of formal taxonomic recognition and is
here newly described as B. zeteticorum Duretto. The remaining two informal taxa are conspecific with B. laxa Duretto.
Additional collections of B. gravicocca Duretto have shown that the taxonomic boundary between it and B. filicifolia
A.Cunn. ex Benth. is not clear and thus B. gravicocca is synonymised under B. filicifolia. Boronia filicifolia was previously
considered to be endemic to the Kimberley region of Western Australia. High levels of endemism of Boronia in the
Northern Territory are discussed. The north-west portion of the Arnhem Land Plateau is determined to be a centre of
diversity and endemism for the genus in the Territory and so is an area of high conservation value. A key to the species
and subspecies of Boronia found in the Northern Territory is provided.

Keywords: Rutaceae, Boronia, Boronia filicifolia, B. gravicocca. B. laxa, B. zeteticorum sp. nov.

INTRODUCTION

Boronia Sm. (Rutaceae) is an Australian genus of
approximately 150 species found in all states and mainland
territories (Wilson 1998; Duretto 1999,2003). The Northern
Territory is a minor centre of diversity for the genus with
17 currently accepted species (12 endemic; one with two
endemic subspecies) (Duretto 1999, 2003; Kerrigan and
Albrecht 2007). In addition, three taxa, B. aff. laxa I
(Northern Plateau, Arnhem Land), B. aff. laxa 2 (Nabarlek,
Arnhem Land) and B. aff. prolixa l (Red Lily Lagoon,
Arnhem Land), were informally recognised by Duretto
and Ladiges (1997) and Duretto (1999). These taxa were
not formally described at the time as there was insufficient
material available and the names have not been adopted in
the literature (e.g. Kerrigan and Albrecht 2007).

Since 1997, a significant number of botanical surveys
have occurred that have resulted in additional material for
many rare species of Boronia , including B. amplectens
Duretto, B. gravicocca Duretto and B. laxa Duretto. This
additional material has allowed several taxonomic problems
to be re-evaluated for the Northern Territory including the
status of the informal taxa. Of the three informal taxa, two,
B. aff. laxa 1 (Northern Plateau, Arnhem Land) and B. aff.
prolixa I (Red Lily Lagoon, Arnhem Land), arc assignable
to B. laxa. Boronia aff. laxa 2 (Nabarlek, Arnhem Land) is
distinct enough to warrant formal taxonomic recognition,
and is formally described below as B. zeteticorum Duretto,

despite being known only from a single collection. Boronia
zeteticorum sp. nov. is placed in B. subseries Grandisepalae
and differs from all other species in that subseries (and B.
subsection Grandisepalae ) by the leaves having distinctly
different indumentum densities on the abaxial and adaxial
surfaces. The species is similar to B. amplectens in that
both have very narrow leaves; a character that could be
considered an apomorphy for the two species and has
proven to be robust with the addition of many collections
of both B. amplectens and B. laxa over the last decade.
Boronia amplectens has stellate hairs with appressed rays
while the stellate hairs of B. zeteticorum are multi-angular
(viz. not appressed).

Boronia gravicocca Duretto was described from a small
number of collections from Bradshaw Station (western
Northern Territory) (Duretto 2003). It was subsequently
targeted for population surveys by the staff of the Northern
Territory Herbarium. Their collections have rendered
B. gravicocca impossible to maintain as distinct from
B. filicifolia A.Cunn. ex Benth. and so the former is here
synonymised under the latter (see below).

MATERIALS AND METHODS

Herbarium material from mainly DNA, but also CANB,
MEL and HO, were studied for this study. Herbarium
abbreviations follow Holmgren et al. (1990).

M. F. Duretto

CLASSIFICATION AND ENDEMISM OF BORONIA
IN THE NORTHERN TERRITORY

Boronia is widely distributed across the ‘Top End’ of the
Northern Territory, which is part of a biogeographical region
that includes north-western Queensland and the Kimberley
region of Western Australia. Boronia is classified into six
sections (Wilson 1998; Duretto 1999, 2003, In press) of
which only B. section Valvatae occurs in north-western
Australia. The species of Boronia in the Northern Territory
are placed in either B. subsection Valvatae ( B. lanceolata F.

Muell., B. rupicola Duretto) or B. subsection Grandisepalae
(remaining 16 species) (Duretto 1999, 2003) (Table 1).

Boronia subsection Valvatae

Within B. subsection Valvatae , B. lanceolata (Northern
Territory, north-western Queensland) is most closely related
to two species from central Queensland in B. series Valvatae
subseries Lanceolatae (Duretto and Ladiges 1999; Duretto
1999, In press). Boronia rupicola (Northern Territory) is
the sole representative of B. series Rupicolae (Duretto
and Ladiges 1999; Duretto 1999). Interestingly, the only
other member of B. subsection Valvatae in north-western
Australia, B. hoipolloi Duretto (north-western Queensland),
is placed in B. series Valvatae subseries Valvatae, and
is closely related to species on Cape York Peninsula
(Queensland) (Duretto and Ladiges 1999; Duretto 1999).

Boronia subsection Valvatae is not found in the Kimberley
region of Western Australia and the three representatives
found in the Northern Territory and north-western
Queensland represent three quite distinct lineages. This
relationship is in contrast to what is found in B. subsection
Grandisepalae.

Table 1. Classification of Boronia in the Northern Territory.

Boronia (all Australian states and mainland territories; c. 150 spp. in 6 sections)

Boronia section Valvatae (W.A., N.T.. Qld, N.S.W., Vic.; 62 spp. in 5 subsections)
subsection Valvatae (N.T., Qld. N.S.W., Vic.; 35 spp. in 4 series)
series Rupicolae (NT.; t sp.)

B. rupicola

series Valvatae (N.T., Qld. N.S.W., Vic.; 24 spp. in 3 subseries)
subseries Lanceolatae (N.T., Qld; 7 spp.)

B. lanceolata (N.T., NW Qld)

subsection Grandisepalae (N W. A., N.T., NW Qld; 20 spp. in 3 series)
series Quadrilatae (N.T.; 2 spp.)

B. cptadrilata, B. viridiflora
series Grandisepalae (N.T.; 8 spp. in 2 subseries)
subseries Verecundae (N.T.; 2 spp.)

B. verecunda, B. xanthastrum
subseries Grandisepalae (N.T.; 6 spp.)

B. amplectens, B. grandisepala (2 subsp.: aciinthopsida and grandisepala), B. laxa, B. prolixa, B. suberosa, B.

zeteticorum sp. nov.

series Lanuginosae (N W.A., N.T., NW Qld; 10 spp. in 3 subseries)
subseries Lanuginosae (N W.A., N.T.. NW Qld; 2 spp.)

B. lanuginosa (NE W.A., N.T.. NW Qld), B. wilsonii (N W.A., NW N.T.)
subseries Jucundae (N W.A., N.T.; 3 spp. )

B. dccumbens (N.T.), B. jucunda (N W.A., NW N.T.), II tolerans (N.T.)
subseries Filicifoliae (N W.A.. NW N.T.; 5 spp.)

B. filicifolia (N W.A., NW N.T.)

Boronia subsection Grandisepalae

Boronia subsection Grandisepalae contains 20 species,
is restricted to north-western Australia (i.e. northern Western
Australia, Northern Territory, north-western Queensland)
and has three scries, Quadrilatae , Grandisepalae and
Lanuginosae (Duretto and Ladiges 1999; Duretto 1999).
The first two series are endemic to the Northern Territory.
Boronia series Quadrilatae (B. quadrilata Duretto,
B. viridiflora Duretto) is restricted to the north-western
Arnhem Land Plateau.

Boronia series Grandisepalae is more widespread and is
classified into two subseries. Boronia subseries Verecundae
(i.e. B. verecunda Duretto, B. xanthastrum Duretto) is
found on the Arnhem Land Plateau and a few outliers to
the west mainly in southern Kakadu National Park though
B. xanthastrum is also found in eastern Arnhem Land.
Boronia subscries Grandisepalae is more widespread,
though only B. grandisepala F. Muell. is found outside the
north-west portion of the Arnhem Land Plateau in eastern
Arnhem Land and south of Mount Brockman to Nitmiluk
National Park and also in the north-western parts of the
Northern Territory. The remaining laxa (i.e. B. amplectens , B.
laxa, B. prolixa Duretto, B. suberosa Duretto, B. zeteticorum
sp. nov.) are narrow endemics of the north-western Arnhem
Land Plateau and nearby outliers, an area that virtually
matches the catchment area of the East Alligator River.

Boronia series Lanuginosae is classified into three
subseries that are all found in the Northern Territory
and the Kimberley Region (northern Western Australia).
Boronia subseries Lanuginosae contains two widespread
species, B. lanuginosa Endl. (Western Australia, Northern
Territory, north-western Queensland) and B. wilsonii

Boronia (Rutaceae) in the Northern Territory

(F. Muell. ex Benth.) Duretto (Western Australia, Northern
Territory), the first species being the only representative of
B. subsection Grandisepalae in north-western Queensland.
Boronia subseries Jucundae contains three species. Two,
B. tolerans Duretto and B. decumbens Duretto, are rare
species of Nitmiluk and Kakadu National Parks. The third
species, B.jucunda Duretto, is found in north-western parts
of the Northern Territory and the eastern Kimberley region.
Boronia subseries Filicifoliae contains five species of the
Kimberley region, one of which, B. filicifolia, extends to
the north-western parts of the Northern Territory.

Of the 18 species of Boronia found in the Northern
Territory, 13 are endemic; eight of these endemics are
restricted to the north-western area of the Arnhem Land
Plateau and nearby outliers roughly within the catchment
area of the East Alligator River. This area measures
approximately 75 x 75 km with the Mount Brockman
outlier in the south-west comer. The remaining endemics are
found to the south of this area on or near the western parts
the Arnhem Land Plateau and outliers south to Nitmiluk
National Park, though B. grandisepala and B. xanthastmm
are found further afield (see above). The distributional
pattern of the species of Boronia found in the Northern
Territory matches that found for all endemic species of the
Northern Territory as presented by Woinarski et al. (2006).
That is, the greatest diversity is found on the north-west
portion of the Arnhem Land Plateau and outliers with the
number of taxa increasing dramatically from south to north
(see also discussion in Duretto and Ladiges 1997). This
area, or rather the catchment area of the Alligator River,
was also identified as a minor hot-spot of endemism in an
Australia-wide study on endemism by Crisp et al. (2001)
and other studies (c.g. Bowman et al. 1988; Dunlop and
Webb 1991). The north-west is typified by many species
with very narrow distributional ranges. In some cases they
do represent aggregates of closely related species.

Boronia subseries Grandisepalae is an excellent
example of an aggregate of many narrow endemics over
a small area with live of its six species restricted to the
north-west plateau area (see above). The four species
with scrambling or lax habit (i.e. B. amplectens, B. laxa,
B. prolixa, B. zeteticorum sp. nov.) form a monophylctic
clade within this group (Duretto and Ladiges 1999: Fig.
10). The relationships of this clade of scrambling species
with B. grandisepala and B. suberosa is equivocal as they
form a trichotomy (see Duretto and Ladiges 1999). As B.
grandisepala is also found outside the region the subseries
may be represented in the area by more than one lineage.

The remaining three species of Boronia endemic to the
north-west Arnhem Plateau area are classified into two
different series, B. series Quadrilatae (with two narrow
endemics) and B. series Rupicolae (monotypic), both of
which are endemic to the region and taxonomically isolated
in their subsections (Duretto and Ladiges 1999; Duretto
1999; Table 1). In a cladistic analysis presented by Duretto
and Ladiges (1999), these series, plus the species in B. series

Quadrilatae , each have a large number of autapomorphies
(see also Duretto 1999) suggesting a long period of isolation
and/or a rapid accumulation of apomorphies.

Also present in the north-west Arnhem Plateau region are
two widespread species, B. lanceolata and B. lanuginosa,
each placed in a different scries to the taxa discussed above
(Table 1). Both species show considerable morphological
variation over their ranges (Duretto 1997, 1999).

Neobyrnesia J.A. Arrnstr., a monotypic genus closely
related to Zieria Sm. (a genus of eastern Australia and New
Caledonia) and Boronia , is also restricted to the north-west
Arnhem Plateau region. Interestingly this genus shares
a number of characters with species of Boronia section
Valvatae, particularly those of B. subseries Grandisepalae,
in the region - the valvatc and persistent petals and similar
seed morphology (Paul G. Wilson pers. comm.; Stace and
Leach 1994; Duretto and Ladiges 1997); the longitudinal
rows on the seed testa are each made of a single line of
tubercles (Armstrong and Powell 1980) as in B. subseries
Grandisepalae and in contrast to Zieria (Duretto and
Ladiges 1997). These characters may indicate a closer
relationship than the current classification suggests or
parallel adaptations to similar niches in the dissected
sandstone landscape. Neobyrnesia suberosa J.A. Arrnstr.
is a cliff specialist, as are B. suberosa (which superficially
resembles N. suberosa as both are shrubs with corky stems
and simple, opposite leaves), B. viridiflora, B. rupicola and
species in other families found in the area (see discussion
under species in Duretto (1999)). Cliff faces are a specialist
niche exploited in a variety of ways by divergent taxa in
the north-west Arnhem Land Plateau region.

Woinarski et al. (2006) reviewed the literature and
discussed several possible mechanisms behind the high
level of endemism found in the north-west region ol the
Arnhem Land Plateau. Within Boronia, the diversity seen
in the region may be the combination of recent species
radiation in conjunction with long-term isolation or refugia.
A recent species radiation, where a large number of closely
related species are found in close proximity, may be what is
seen in B. subseries Grandisepalae. The endemic series (and
Neobyrnesia ) probably also represent autochthonous taxa
that have been in the region and/or isolated for a longer time.
In effect, for these taxa, the region has acted as a refugium.
In addition, the region is also home to widespread species
that may be relatively recent additions to the local flora.

When the species of Boronia found in the north-west
portion of the Arnhem Land Plateau are highlighted on the
cladogram of/?, section Valvatae presented by Duretto and
Ladiges (1999: Figs 9 and 10) at least five distinct lineages
are identified: B. lanceolata, B. lanuginosa, B. rupicola, B.
scries Quadrilatae, B. subscrics Grandisepalae (see above).
Only two of these lineages are represented by more than one
species. The area is not only the most diverse area of the
Northern Territory (and north-western Australia) in terms
of the number of species of Boronia, but it is also the most
diverse in terms of the number of infrageneric taxa present.

M. F. Duretto

Thus the north-west portion of the Arnhem Land Plateau
(i.e. the catchment area of the East Alligator River) is an
area of extremely high conservation value.

TAXONOMY

Boronia filicifolia A.Cunn. ex Benth., FI. Austral. 1:

311 (1863)

Type: Australia, Western Australia, Montague and York
Sounds, N.W. Australia, 1820, A.Cunningham 220. third
voyage of the “Mermaid” (lectotype. /zV/e Duretto Nuytsia
11: 332 (1997): K n.v. (photograph AD, cibachrome MEL);
isolectotype: BM n.v. (transparency MEL, photograph
PERTH).

Synonymy.

Boronia gravicocca Duretto (2003): 123, syn. nov.

Illustrations. Wheeler et al., (1992) FI. Kimberley
Region 669, Figs 206 A1 and A2; Duretto (2003) Muelleria
17: 102, figs 12 K-M, as B. gravicocca; Duretto (2005)
Australian Plants 23 (183): 90 (2005).

Remarks. For a full description on this species see
Duretto (1997, 1999, 2003).

Distribution and ecology. Boronia filicifolia occurs in
the Mitchell River and Port Warrender areas (north-west
Kimberley region. Western Australia) and disjunctly at
Bradshaw Station (Victoria River area, Northern Territory).
It is found in heath and open woodland on sandstones and
quartzites. Flowering December-June; fruiting (December,
Feb.) June-July.

Boronia laxa Duretto, Austral. Syst. Bot. 10: 279
(1997)

Type. Australia, Northern Territory, Site FF, c. 30 km
SE of Jabiru, 12°55’S 132°59’E, 30 March 1981, L.A.
Craven 6600 (holotype: CANB 338123 (photograph HO);
isotypes: AD, DNA 20968 (transparency MEL), MEL
2041245, P, US).

Synonymy.

Boronia aff. laxa l (Northern Plateau, Arnhem Land)
Duretto and Ladiges (1997): 282.

Boronia atf. prolixa 1 (Red Lily Lagoon, Arnhem Land),
Duretto and Ladiges (1997): 285.

Illustration. Duretto and Ladiges (1997), Austral. Svst.
Bot. 10: 280, fig. 20 a-b.

Remarks. For full descriptions as well as ecological and
distributional data on this species see Duretto and Ladiges
(1997) and Duretto (1999).

Boronia zeteticorum Duretto, sp. nov.

(Fig. 1)

A B. laxa Duretto foliis angustioribus (1.5-3.5, non
2.5-10 mm latis) indumenta abaxiali denso, adaxiali
moderate denso dijfert.

Differs from B. laxa by its narrower leaves (1.5-3.5 mm
wide, not 2.5-10 mm wide) with a dense indumentum on

the abaxial surface and a moderately dense indumentum
on the adaxial surface.

Type. Australia, Northern Territory, Nabarlek, Arnhem
Land, 12°19’S 133°19’E, 23 March 1989, R. Hinz 467
(holotype: DNA 43905 (transparency MEL); isotype:
CANB).

Synonymy.

Boronia aff. laxa 2 (Nabarlek, Arnhem Land) Duretto
and Ladiges 1997: 282.

Illustration. Duretto and Ladiges, Austral. Syst. Bot.
10. 270, fig. 16i-j (leaf hairs); 276, fig. 19i (seed surface)
(1997), all as B. sp. aff. laxa 2.

Specimen examined. Known from the type material
only.

Description. Semi-prostrate, much branched subshrub
to 50 cm long; multi-angular stellate hairs sessile, 6-25+
rays per hair; rays unicellular, unfused, firm, straight,
glossy, smooth, yellow-white, 0.1-0.2 mm long. Branches
not obviously glandular, with little cork development,
with a moderately dense stellate indumentum. Leaves
simple; petiole 0.5-1.5 mm long; lamina 10-35 mm
long, 1.5-3.5 mm wide, narrowly elliptic, tip acute, base
attenuate, margins fiat, slightly discolorous, paler beneath,
not obviously to slightly glandular, midrib raised abaxially,
slightly impressed adaxially; adaxial surface with moderately
dense stellate indumentum (Duretto and Ladiges 1997:
Fig 16i); abaxial surface with dense stellate indumentum
(Duretto and Ladiges 1997: Fig 16j). Inflorescence axillary,
1-flowered, with a dense stellate indumentum; peduncle
0.5 mm long; prophylls to 2 mm long and 0.5 mm wide;
metaxyphylls minute to 1 mm long; pedicels c. 1.5 mm long.
Sepals white, longer and wider than petals, 3.5-4 mm long,
c. 2 mm wide, enlarging to 6 mm long and 3.5 mm wide
with fruit, ovate to dcltate, tip acute to acuminate, valvate
in bud, persistent with mature fruit; adaxial surface with
dense and minute indumentum along margins, becoming
glabrous towards centre and base; abaxial surface with
moderately dense stellate indumentum. Petals white,
3-3.5 mm long, 1-1.5 mm wide, enlarging to 4.5-5 mm
long in fruit, valvate in bud, persistent with mature fruit;
adaxial surface with sparse stellate indumentum becoming
glabrous towards base; abaxial surface with moderately
dense to dense stellate indumentum. Stamens: filaments
bearing stiff, simple hairs abaxially and on margins below
glandular tip; antesepalous filaments clavate and suddenly
narrowing to anther connective, 1.5 mm long, distal 0.75
mm prominently glandular; antepetalous filaments slightly
glandular, c. 1 mm long; antepetalous anthers much
larger than antesepalous anthers. Ovary glabrous; style
glabrous. Cocci with moderately dense simple and stellate
indumentum, c. 4 mm long, c. 2 mm wide. Seeds black,
shining, c. 3 mm long, 1.5 mm wide, elliptical with adaxial
side flattened and with prominent ridge; testa striated, at
magnification tuberculate to colliculate with units fused
to form longitudinal ridges 33-50 pm apart; ridge units

Boronia (Rutaceae) in the Northern Territory

DNA 43908

"Boronitc. Co r por.-tf'o

Dim HOCOT

HF r. f-A, -u.p

Tasmanian herbarium ihoi

NATIONAL HERBARIUM OF VICTORIA

j! fine- jLrv$? '2—

■ [let? Tl-I L-A'y, ^-‘7. Svrr. ier.
It?: ?n f ' C '“'V.3
Dcterminavit /^/ ^ /Jk

Seen for

f/ora of Australia

northern territory herbarium

DARWIN (DNA). AUSTRALIA

rlek, Arnhca Land.

12.19S 133.19E

E/DIST. t HT/DC

jtonc country) rambling to protrate
with white flowcrn.

- , •■»!-> tOflQ

rutackak

Fig. 1. Holotype of Boronia zeteticorum sp. nov. (Hinz 467 , DNA 43905). Illustration 55% actual size.

M. F. Duretto

unicellular, smooth, anticlinal walls more or less visible,
30-43 jam across (Duretto and Ladiges 1997: Fig 19i).

Distribution and ecology. Boroniazeteticorum is known
only from the Nabarlek area (Northern Territory) (Duretto
and Ladiges 1997: Fig. 2; or Duretto 1999: Fig 16). The
only ecological information given with the single collection
is that it was made in sandstone country. The material was
collected in March and has both flowers and fruit.

Remarks. Boronia zeteticorum differs from other
species in B. subsection Grandisepalae by the surfaces
of the leaves having contrasting indumentum densities,
i.e. a dense indumentum on the abaxial surface but a
moderately dense indumentum on the adaxial surface
(Duretto and Ladiges 1997: Fig. 16ij). It is placed in B.
subseries Grandisepalae and is semiprostrate with weak
stems, as are B. amplectens, B. laxa and B. prolixa with
which it is probably forms a monophyletic group. It differs
from B. laxa and B. prolixa by its very narrow leaves: a
character it shares with B. amplectens. Boronia amplectens
has stellate hairs with rays apressed to the surface while the
stellate hairs of B. zeteticorum are multi-angular.

Conservation Status. The species is known from a
single collection made in 1989. Field surveys are required
urgently to ascertain the correct conservation status for
this species.

Etymology. The specific epithet is derived from the
Greek, zetetikos (disposed to search), as coined by Craven
and Jones (1991: 529), and ‘honours those persons who for
their enjoyment explore natural vegetation communities to
become familiar with, and collect, their constituent species’.
The systematic study of Boronia , like many genera of
plants, would not be where it is today without the collections
made by such people in remote places.

KEY TO SPECIES AND SUBSPECIES OF BORONIA
FOUND IN THE NORTHERN TERRITORY

For descriptions, ecology and distributional data see
Duretto (1999); for B. filicifoliae see also Duretto (2003)
to include information given under B. gravicocca.

la. Sepals much shorter than petals (< 'A length);
antepetalous anthers c. equal to antesepalous

anthers.2

1 b. Sepals nearly as long (> 'A) to much longer than petals;
antepetalous anthers much larger than antesepalous
anthers.....3

2a. Leaves simple, midrib on the abaxial surface
prominently raised; erect or very rarely pendulous
shrubs; petals 2-7 mm long; staminal filaments

glabrous or with 1-3 hairs. B. lanceolata

2b. Leaves 1-7-folilate, midrib of abaxial surface not
prominently raised; pendulous shrubs growing on
cliff faces; petals 2-3 mm long; staminal filaments
hirsute. B. rupicola

3a. Leaves pinnate.4

3b. Leaves simple.9

4a. Sepals shorter and narrower than petals, 2-3.5 mm
long; at least some leaves 27—41 -loliolate, with a
sparse to moderately dense indumentum (epidermis
clearly visible); lateral leaflets c. rhombic to circular

to broadly elliptic; petiole to 2 mm long.

. B. filicifolia

4b. Sepals longer and wider than petals, 3-15 mm long;
leaves 3-27(-35)-foliolate (if > 30 leaflets then some
petioles > 3 mm long), glabrous to densely stellate
tomentose; lateral leaflets linear to narrow-elliptic,
rarely elliptic; petiole to 7 mm long.5

5a. Leaves petiolate, though petiole sometimes as
small as c. 0.5 mm long; leaflets linear-elliptic to
elliptic, the margins plane or recurved to revolute;
lamina glabrescent or with a sparse to dense, stellate

indumentum. 6

5b. Leaves sessile; leaflets linear-elliptic, plane or slightly
recurved along margins; lamina glabrescent or with a
sparse, stellate indumentum.7

6a. Leaflets linear to narrowly elliptic, so revolute that
abaxial surface not usually visible; sepals (4—)5—14
mm long, glabrous or with a sparse to dense

indumentum (Top End). B. lanuginosa

6b. Leaflets elliptic to lanceolate, abaxial surface visible;
sepals 5-9 mm long, with a dense indumentum
(Victoria River area). B. wilsonii

7a. Branches obviously glandular; leaves 3-foliolate.

. B. jucunda

7b. Branches eglandular; leaves (3-)5-9-foliolate.8

8a. Plants decumbent, with a sparse to moderately dense,
simple indumentum, stellate hairs rare; leaflet-margins

slightly recurved. B. decumbens

8b. Plants erect, with a sparse stellate indumentum;
leaflets plane. B. tolerans

9a. Plants, other than flowers, glabrous, glaucous
(especially leaves); stems distinctly quadrangular, at

least on young shoots...10

9b. Plants sparsely to densely hirsute, not glaucous; stems
terete to slightly quadrangular.11

10a. Erect shrub growing on ridges; leaves sessile; sepals
6-13 mm long; petals 4-5 mm long....5. quadrilata
10b. Horizontal shrub growing from cliff faces; leaves

petiolate; sepals and petals 2.5-3 mm long.

. B. viridiflora

Boronia (Rutaceae) in the Northern Territory

11a. Stellate hairs prominently stalked, rays 0.5-1 mm

long; cocci glabrous.12

lib. Stellate hairs without prominent stalks, rays to 0.5
mm long; cocci hirsute.13

12a. Hairs white and flexuous, new shoots pinkish to
white; leaves narrowly elliptic; adaxial surface of

petal glabrous or hirsute. B. verecunda

12b. Hairs yellow and straight, new shoots yellow; leaves

elliptic; adaxial surface of petal hirsute.

... B. xanthastrum

13a. Older stems with massively developed cork;
indumentum of leaves usually difficult to see with
the unaided eye, rays of hairs to 0.1-0.3 mm long...

... B. suberosa

13b. Older stems not corky; indumentum of leaves clearly
visible to the unaided eye, rays of hairs 0.1-0.5 mm
long.14

14a. Plants erect (rarely sprawling but then with a hoary,
dense indumentum), with a moderately dense to
dense indumentum; sepals greater than 7 mm long at
anthesis (rarely less than 7 mm long but then plant

with a hoary, dense indumentum).15

14b. Plants sprawling, sparsely to moderately hirsute
(rarely densely hirsute on the abaxial leaf-surface
only); sepals less than 8 mm long at anthesis,
sometimes enlarging to 11 mm long as fruit matures
.16

15a. Leaves with a very dense indumentum, epidermis not

visible. B. grandisepala subsp. grandisepala

15b. Leaves with a moderately dense indumentum, leaf

epidermis visible.

. B. grandisepala subsp. acanthopsida

16a. Leaves narrow-elliptic, 1-4 mm wide.17

16b. Leaves elliptic, lanceolate, ovate, > 5 mm wide... 18

17a. Leaves: indumentum of both abaxial and adaxial

surfaces similar, sparse; rays of hairs appressed.

. B. amplectens

17b. Leaves: indumentum of abaxial and adaxial surfaces
markedly different; that of the abaxial surface dense;
that of the adaxial surface moderately dense; hairs
multi-angular, rays not appressed. B. zeteticomm

18a. At anthesis Dowers stalks (peduncle and pedicel) 2-7
mm long, not bent at prophylls (bracts); leaves elliptic

to lanceolate. B. laxa

18b. At anthesis at least some flowers stalks (peduncle
and pedicel) > 10 mm long, often bent at prophylls
(bracts); leaves ovate to lanceolate. B. prolixa

ACKNOWLEDGMENTS

I would like to thank the Director and staff of DNA

for access to the collections and rapid processing of loan

material as well as Neville Walsh (MEL) for the Latin

diagnosis.

REFERENCES

Armstrong, J.A. and Powell, J.J.M. 1980. Neobyrnesia (Rutaceae),
a new genus endemic to northern Australia. Telopea 1:
399-408.

Bowman, D.M.J.S.. Wilson, B.A. and Dunlop, C.R. 1988. Preliminary
biogeographic analysis of the Northern Territory vascular flora.
Australian Journal of Botany 36: 503-517.

Craven, L.A. and Jones, S.R. 1991. A taxonomic review of
Homoranthus and two new species of Darwinia (both
Myrtaceac, Chamelaucieae). Australian Systematic Botany
4:513-533.

Crisp, M.D., Laffan, S., Linder, H.P. and Monro, A. 2001. Endemism
in the Australian flora. Journal of Biogeography 28: 183—
198.

Dunlop, C.R. and Webb, L.J. 1991. Flora and vegetation. Pp. 41-60
In: C.D. Haynes, C.D., Ridpath, M.G. and Williams, M.A.J.
(eds) Monsoonal Australia: landscape, ecology and man in
northern lowlands. A.A. Balkema: Roterdam.

Duretto, M.F. 1997. Taxonomic notes on Boronia species of
north-western Australia, including a revision of the Boronia
lanuginosa group (Boronia section Valvatae: Rutaceae).
Nuytsia 11: 301-346.

Duretto, M.F. 1999. Systematics of Boronia section Valvatae sensu
lato (Rutaceae). Muelleria 12: 1-131.

Duretto, M.F. 2003. Notes on Boronia (Rutaceae) in eastern and
northern Australia. Muelleria 17: 19-135.

Duretto, M.F. In press. Description of a new subsection and two
new subseries for Boronia section Valvatae (Rutaceae).
Austrobaileya.

Duretto. M.F. and Ladiges, P.Y. 1997. Morphological variation within
the Boronia grandisepala group (Rutaceae) and the description
of nine taxa endemic to the Northern Territory, Australia.
Australian Systematic Botany 10: 249-302.

Duretto, M.F. and Ladiges, P.Y. 1999. A cladistic analysis of Boronia
section Valvatae (Rutaceae). Australian Systematic Botany
11:636-665.

Holmgren, P.K., Holmgren, N.ll. and Barnett, L. 1990. Index
Herbariorum. Part /. The Herbarium of the World, 8th edn.
New York Botanical Gardens: New York.

Kerrigan, R.A. and Albrecht, D.E. 2007. Checklist of NT vascular
plant species. Northern Territory' Herbarium: Darwin.

Stace, H.M. and Leach, G.J. 1994. Cytological notes in Rutaceae. 2:
Neobyrnesia suberosa. Telopea 6: 167-168.

Wilson, P.G. 1998. New names and new taxa in the genus Boronia
(Rutaceae) from Western Australia, with notes on seed
characters. Nuytsia 12: 119-154.

Woinarski, J.C.Z., Hempel, C., Cowie, I.,Brennan, K., Kerrigan, R.,
Leach, G. and Russcll-Smilh, J. 2006. Distributional pattern
of plant species endemic to the Northern Territory, Australia.
Australian Journal of Botany 54: 627-640.

Accepted 26 August 2008

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 15-21

Jaw growth and replacement in Diopatva aciculata (Annelida: Onuphidae)

HANNELORE PAXTON 1 AND MILADA SAFARIK 2

'Department of Biological Sciences, Macquarie University, Sydney, NSW 2109, AUSTRALIA
Email: hpaxton@rna.bio.mq.edit.au
2 Aquabait Pty Ltd, PO Box 5107, Dora Creek, NSW2264, AUSTRALIA

ABSTRACT

Jaw development was studied in 1 - to 45-day old Diopatra aciculata (fertilisation to 44-chaetigers). Both mandibles and
maxillae became visible in 4- to 6-chaetiger worms. The initial protomandibles become embedded in the adult mandibles.
The larval maxillae consist of a left fang-like and right small serrated clement, a pair of serrated plates, distal ridges, and
a single carrier. Larval maxillae were replaced by juvenile maxillae in 14- to 15-chaetiger worms. Juvenile maxillae have
adult-like carriers. Mil and MIV, but MI are proximally dentate with a distal hook; Mill and MV are absent. First adult
maxillae were seen in 21 -chaetiger worms. Each subsequent moult results in larger but otherwise identical apparatuses.
The moult increment (difference in size between successive moults) was established as 1.34 and used to extrapolate
estimates of size classes. Based on these calculations our largest wonns had their fourth set of adult maxillae, having
moulted the adult maxillae every three to four days.

Keywords: Polychacta, Eunicida, jaw apparatus, mandibles, maxillae, size classes, moulting.

INTRODUCTION

The tubicolous onuphid polychaete Diopatra acicu¬
lata Knox and Cameron. 1971 occurs in shallow depths
along the southern shores of Australia (Paxton 1993). It is
collected as prized fishing bait and commercially farmed
as bait and food in the conditioning of prawn broodstock
(Safarik et al. 2006). The worms have a lifespan of about
five years (Safarik et al. 2006) and are dioecious. Mature
eggs are about 230 pm in diameter. After a free-swimming
lecithotrophic larval stage of 4 to 6 days D. aciculata settles,
starts to build its tube and feed.

As typical for Eunicida, D. aciculata has a complex
jaw apparatus consisting of ventral mandibles and dorsal
maxillae. While mandibles grow during the lifetime of
an individual, the maxillae arc moulted or replaced many
times. Larval and juvenile jaws have been reported for
several species of Onuphidae (Table 1), although most of
these reports were anecdotal. The most complete study is
of Kinbergonuphis simoni Santos et al., 1981, following the
development of larval to adult maxillae and confirming the
replacement of larv al maxillae by adult types (Hsieh and
Simon 1987), previously only known for Mooreonuphis
jonesi Fauchald, 1982 (Fauchald 1982).

Shedding and replacement of adult maxillae in
D. aciculata, comparable to arthropod moulting, has been
reported (Paxton 2005). Replacement of maxillary jaws in
Eunicida has been recently reviewed, suggesting that the
type of moulting observed in Onuphidae may also take place
in Eunicidae and Lumbrineridae (Paxton 2006). However,
it is not known how often the maxillae are replaced. While
those of older adult worms may moult infrequently or not

at all, it is expected that those of young worms undergo
frequent moults to keep pace with the rapidly growing
body.

The aim of the present paper was to study the de¬
velopment of jaws in very young (1 to 45 days old) D.
aciculata in order to describe and illustrate the growth of
the mandibles and replacement of maxillae. It was hoped
that relating the size of the maxillary elements to the age
and size of the worms might reveal age classes and thus a
possible moult cycle.

MATERIALS AND METHODS

Diopatra aciculata were reared in outside ponds at
the Aquabait Pty. Ltd. aquaculture facility at Dora Creek,
NSW, Australia. The study material ranged from newly
fertilised eggs (day 1) to animals consisting of44 chaetigers
(maximum of 45 days). The three stages of the maxillary
apparatus are referred to as larval, juvenile and adult, even
though they may not correspond to the state of maturity of
the animals. Animals were mounted on a slide with diluted
glycerine added to the edge of the coverslip and examined
with a compound microscope. The jaws were studied in
situ. The number of chaetigers was used as an expression
of body size. The elements of the adult maxillary apparatus
are numbered in the conventional way in Roman numerals
progressing from posterior to anterior, so that for instance
MIL refers to the left most posterior maxilla, following the
maxillary carrier (Fig. 1A). The length of maxilla 1 was
taken as an indicator of the size of the complete maxillary
apparatus. The maxilla I was drawn with the aid of a camera
lucida and measured on the drawing.

H. Paxton and M. Safarik

Fig. 1. Jaws of Diopatra aciculata: A, adult maxillae of 42-chactigcr specimen, dorsal view; B, protomandibles of 4-chaetiger specimen,
dorsal view; C, protomandibles of 5-chaetiger specimen, dorsal view; D. mandibles of 42-chactiger specimen, showing incorporated
protomandibles, ventral view; E, developing larval maxillae in 4-chaetiger specimen, dorsal view; F, larval maxillae with short carrier in
4-chaetiger specimen, dorsal view; G, carrier of 5-chaetiger specimen, dorsal view; H, same of 12-chaetiger specimen, dorsal view; 1, juvenile
maxillae of 19-chaetiger specimen.

Jaw growth and replacement in Diopatra aciculata

The moult increment is a term used in entomology
to describe the increase in size between two successive
moults. Dividing the postmoult size by the premoult size
is a constant for a certain species and represents the moult
increment (Gullan and Cranston 1994).

RESULTS

Overall growth. The overall growth of the worms as
expressed in numbers of chaetigers over time is shown in
Figure 2. The largest animals attained a size of 44 chacti-
gers. We are aware of the shortcomings of the quantitative
nature of this data resulting from some time periods when
sampling was not possible. Due to the scarcity of the mate¬
rial we have combined the results from two separate cohorts.
While this has bolstered the numbers, it has introduced a
greater range of variation due to the different growth rates
of the two cohorts. I lowever, in the absence of any previous
study of this kind, we feel that the qualitative nature of the
data overrides the quantitative problems.

Jaw growth - mandibles. The anterior part of the cut¬
ting plate is the first part of the mandibles to become visible
in 4-chaetiger worms, when they are four to six days old.
The plates initially appear as lightly sclerotised structures
(Fig. 1B) and become H-like as more material is deposited
(Fig. 1C). With continuous growth they develop the typi¬
cal adult form which has proximal long shafts and a distal
cutting plate (Fig. 1D).

Jaw replacement - maxillae. The maxillary apparatus
becomes visible at the same time as the mandibles. The
appearance and moulting sequence of the maxillae are
summarised in Table 2.

The tips of the teeth or serrations are the first parts of the
larval maxillae to become visible (Fig. 1E) and within a few
days the whole apparatus is darkly sclerotised (Fig. 1F). It
consists of a single narrow carrier, a large serrated left fang¬
like element, a very small serrated right element, a large
pair of serrated plates, and a distal pair of ridges. The initial
carrier is a short cup-shaped structure that articulates with
the left fang-like element, has a small extension articulating
with the right small plate, and a short posterior extension.
This extension rapidly grows and appears to form a tube¬
like structure (Fig. 1G), that in some preparations opens
posteriorly, displaying growth lines (Fig. 1 FI).

The larval maxillae are shed, swallowed, and replaced
by the juvenile maxillae (Fig. II) in 14- to 15-chaetiger
worms. That apparatus is weakly sclerotised, particularly
the distal part. The juvenile maxillae consist of carriers,
paired asymmetrical MI that are proximally denticulated
and have a distal hook, paired denticulated Mil and less
distinct paired MIV. Maxillae III and V appearto be absent.
The first adult maxillae (Fig. I A) appear in 21-chaetiger
worms, consisting of the typical adult form. Subsequent
moults result in larger but otherwise identical maxillae.

Size classes. The length of maxilla 1 was taken as an
indicator of the size of the total maxillary apparatus and
was found to range from 80-105 pm for juvenile and

o 5 10 15 20 25 30 35 40 45 50

Age in days

Fig. 2. Growth curve of Diopatra aciculata indicating stage of maxillae of each specimen, as shown in key in lower right: (L) larval-
(J) juvenile; (A) adult stage A; (B) adult stage B; (C) adult stage C; (D) adult stage D.

H. Paxton and M. Safarik

Fig. 3. Frequency histogram of MI length of juvenile (J) and adult maxillae of stages (AD) o (Diopatra aciculata. The mean length of each
stage is indicated by asterisk and value above the respective column.

120-310 pm for adult jaws. A frequency histogram of the
length of maxilla I was prepared to reveal size classes (Fig.
4). It demonstrated a well defined juvenile class. The other
classes appear less clear, partly due to lack of material (class
A) and continuity between classes B and C.

In the absence of clear size classes we endeavoured
to discover the moult cycle utilising the moult increment
(Table 3). The known moult increments came from two

individuals with recently moulted maxillae in situ and
discarded old sets recovered from the gut. A 31-chaetiger
specimen (35 days old) had a moult increment of 1.32, while
an 89-chaetiger specimen (82 days old) from a different
study had a moult increment of 1.38. Assuming that the
moult increment between the juvenile and adult maxillae
equals that of consecutive adult maxillae, we calculated the
value by utilising the mean size of four earliest adult and

Table 1. Reports of protomandibles, larval and juvenile maxillae for the family Onuphidae.

Species

Protomandiblcs

Size of norm

Larval maxillae
Size of norm

Juvenile maxillae
Size of norm

Reference

Diopatra aciculata Knox and
Cameron, 1971

4- to 20-chaetiger

same

14- to 28-chaetiger

present study

Diopatra cuprea Bose, 1802

5- to 6-chaetiger

same

Allen 1959

Diopatra lilliputiana Paxton, 1993

15-chaetiger

same

Paxton 1993

Diopatra cf. marocensis Paxton et
at., 1995 as D. cuprea

11-chaetiger

same

Monro 1924

Hyalinoecia araucana Carrasco,
1983

13-chaetiger

same

Carrasco 1983

Hyalinoecia incubans Orensanz,
1990

7-chaetiger

same

Orensanz 1990

Kinbergonuphis simoni (Santos et
al, 1981)

5- to 14-chaetiger

same

15-chaetiger

Hsieh and Simon 1987

Leptoecia abyssorum Chamberlin,

1919 as Paronuphis abyssorum

9-chaetiger

same

Averincev 1972

Mooreonuphis jonesi Fauchald,

1982

8-chaetiger

13-chaetiger

Fauchald 1982

Onuphis elegans (Johnson, 1901)

5- and 7-chaetiger

5-chaetiger

14-chaetiger

Blake 1975

Jaw growth and replacement in Diopatra aciculata

Table 2. Appearance and moulting sequence of maxillary apparatus of D. aciculata.

Type of maxillae

Larval (Fig. IF)

Juvenile (Fig. II)

Adult (Fig. 1 A)

Number and type of elements
in posterior to anterior order

Single carrier

Left fang-like element; small right serrated element
(?M1)

Pair of large serrated plates (?M1I)

Pair of ridges (?MIV)

Paired carriers

Paired asymmetrical MI, denticulate with distal
hook

Paired denticulated Mil
Paired M1V
Paired carriers
Paired falcate MI
Paired denticulate Mil
Single left denticulate Mill
Paired MIV
Paired MV

Age and size of worm

Days since fertilisation Number of chaetigers

4 to 6 4 to 6

14 or 15

21 to 29

ten juvenile MI, establishing the value as 1.33. Thus, the
available data indicate a mean moult increment of 1.34.

To obtain an estimate of size classes we used the moult
increment lor extrapolation. The smallest and largest ju¬
venile MI values were multiplied by the moult increment
to obtain the range of the adult (A) size class. Three more
size classes were extrapolated in the same manner to give
the ranges for four size classes (Table 4). The estimated
size classes have been indicated for each individual in
the growth curve (Fig. 2), demonstrating that the first and
second moults occurred more or less at a certain chaetiger
number but that the later stages overlapped. The estimated
size classes were superimposed on the frequency histogram
of MI length (Fig. 3), indicating again a well defined juve¬
nile class but indistinct adult size classes.

DISCUSSION

It has been stated that the mandibles appear before
the maxillae in the labidognath taxa (Kielan-Jaworowska
1966). This is not the case for D. aciculata and most likely
other labidognath taxa as well.

While larval mandibles have been illustrated in several
studies (Table 1), their relationship to the adult mandibles
is generally not fully appreciated. The sclerotiscd larval
mandibles are here referred to as protomandibles and can
be noted as a dark, X-shapcd structure through the body
wall (Fig. 1C). They will be enlarged by the exterior depo¬
sition of sclerotiscd proteins and carbonates, throughout
life, at the areas in contact with the cuticular epithelium,
i.c. ventrally and laterally. The protomandibles arc visible
in young worms in a ventral view (Fig. 1D) and sometimes
still visible in adults as small dark lines. In decalcified
mandibles the protomandibles are sometimes longer than
the sclerotiscd plate and have been referred to as ‘Stacheln’
or thorns by von 1 latTner (1959).

Larval maxillae are known for a number of species
belonging to several genera of Onuphidae (Table 1). The
main elements of the larval maxillae of all but one species
were reported to consist of proximal narrow carriers, a pair
of plates, a single left fang-like element and a distal pair
of ridges. The only exception, Onuphis elegans (Johnston
1901), was reported as having paired fang-like elements
(Blake 1975). The otherwise remarkable similarity of the

Table 3. Calculation of moult increment of maxillary apparatus of D. aciculata. *) specimen from another study (Paxton 2006)

Method

Number

of worms

Stage of maxillae

Size of worms

Size of premoult Ml

Size of postmoult MI

Moult

increment

Observation

early adult

31-chaetiger

195 pm

1.32

Observation

adult

89-chaetiger*

840 pm

1160 pm

1.38

Estimate

earliest juv./

14- to 21-chae-

80-105, x=93 pm

120-125, x=124

1.33

adults

tiger

Mean of 3 values =

1.34

H. Paxton and M. Safarik

larval maxillary elements of O. elegans to those of the other
species needs investigation.

The larval maxillae of D. aciculata (Fig. 1F) differ from
those reported for other onuphids in having an additional
very small serrated right element. This element is difficult
to observe and presumably has been overlooked in all but
one report. Krohn and Schneider (1867) reported several
annelid larvae. One of these, observed at Madeira, was
followed to the 5-chaetiger stage when it possessed five
rudimentary prostomial appendages and the larval jaw
apparatus. The authors concluded that the jaws indicated
that it belonged to the family Eunicidae and because of the
number of ‘antennae’ probably to the genus Eunice. We
now know that Eunice at that stage has only one median
antenna, and that the larva must have been an onuphid.
The amazing part of the description and small drawing of
the jaw apparatus is that Krohn observed the small right
maxillary element that has presumably been overlooked
by all subsequent researchers.

The larval maxillae are replaced by the juvenile maxillae
(Fig. 11). These resemble the adult apparatus with short,
broad carriers, paired forceps-like Ml and paired Mil, and
were referred to as adult jaws by Hsieh and Simon (1987).
The juvenile MI differs remarkably from the adult type in
that it is proximally medially dentate with a distal hook.
The right hook is longer than the left one, and is similar to
the prionognath type MI of the Oenonidae (e.g. Arabella).
The apparatus is weakly sclerotised, particularly the distal
part. In our specimens MI and Mil were clearly visible,
and MIV weakly visible. The single left Mill and paired
MV of the adult maxillae appeared to be absent. However,
Fauchald (1982) described and figured Mill but not MV
for Mooreonuphis jonesi while Hsieh and Simon reported
MV but not Mill for Kinbeigonupliis simoni. We think it is
probable that the Mill is not present in the juvenile apparatus
and makes its first appearance in the adult apparatus. The
presumed homology of the larval and juvenile maxillary
elements and their evolutionary significance will be
discussed elsewhere (Paxton in press).

Once the juvenile maxillae were replaced by the adult
type, each additional moult resulted in slightly larger
but otherwise identical maxillae. Figure 3 is a frequency
histogram of maxilla I in an attempt to demonstrate size
classes. A clear juvenile class was demonstrated while the
other classes were inconclusive. This is not only a result of
the scarcity of the data. A similar attempt to estimate size
classes of the fossil polychaete Kettnerites ( Kettnerites )
bankvaetensis Bergman, 1987, was carried out by Bergman
(1989). He measured 134 jaws (MI) ranging in size from
200 pm to 890 pm and did not obtain distinct size classes.
Bergman discussed some factors affecting a polymodal size
frequency distribution of Eunicida jaws and re (erred to a
statement by P.J.W. Olive that with polychaetcs it is virtu¬
ally impossible to identify size classes beyond the second
class (P.J.W. Olive pers. comm, to C. Bergman, 1986). The

Table 4. Estimated size classes of adult maxilla 1 of D. aciculata.

Size class

Range of size (pm)

Mean (pm)

Adult A

107-141

124

Adult B

144-189

167

Adult C

192-253

223

Adult D

258-339

299

larger classes tended to merge due to individual growth
rates, food supply and genetics.

The difference in size between two instars in insects
is called the moult increment. The increase of sclerotised
parts, such as the head capsule, a regular linear progression
in successive instars, is known as Dyar’s rule (Gullan and
Cranston 1994). Our moult increment was established as
1.34 (Table 3) and is comparable to the usual insect value
of 1.4 (Hinton and Mackerras 1970). Using this index,
we have estimated size classes of adult maxillae (Table
4) based on the range of juvenile jaws, indicated these on
Figure 3 and marked the points on Figure 2 according to
the extrapolated size classes. During the 45 days of the
experiment, the maxillae retained for the longest period
were the larval maxillae. The worms consisted of 14-15
chaetigers (28-35 days old) at the moult to the juvenile jaws.
This timing is identical with the corresponding moult in the
brooding A-/, jonesi and K. simoni. Hsieh and Simon (1987)
linked this event with leaving the maternal tube to construct
tubes on their own and the commencement of feeding. Our
D. aciculata, lecithotrophic only fora brief free-swimming
larval stage, settles, starts to construct its lube and to feed
at the age of 4-6 days (5-chaetigers) as evidenced by fecal
pellets in the gut. This indicates that the timing of the first
moult, i.e. larval to juvenile maxillae, is not governed by
ecological but developmental clues.

Based on these estimations (Figs 2,3), our largest wonns
(42^14 chaetigers) had their fourth set of adult maxillae,
having undergone a moulting process every 3-4 days
since attaining the juvenile maxillae. As the growth rate
slows down, the frequency of moulting presumably slows
down. Specimens from a different study (Safarik et al.
2006) that were raised for seven months and consisted of
150-186 chaetigers, had maxillae I of 1.6-1.9 mm length,
indicating that these were their tenth adult maxillae, leading
to the conclusion that the rate of moulting slows down
dramatically with time, unless it occurs with less or no size
increase in later life.

ACKNOWLEDGMENTS

We are grateful to Les Safarik for allowing us to use
the facilities of Aquabait Pty. Ltd. for this study. We also
thank Gunter Purschke and Mats Eriksson for review of the
manuscript and valuable suggestions.

Jaw growth and replacement in Diopatra aciculata

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Gullan, P.J. and Cranston, P.S. 1994. The insects: an outline of
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Hinton, 11.E. and Mackerras, M.J. 1970. Reproduction and
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XXL Antarctic Research Series 52: 1-183.

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Onuphidae) from Australia, with a discussion of developmental
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Accepted 26 August 2008

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 23-31

Feeding selectivity of sesarmid crabs from northern Australian mangrove forests

CHANDRA P. SALGADO KENT* AND KEITH A. McGUINNESS

School of Science and Primary Industries, Faculty of Education, Health & Science, Charles Darwin University,

Darwin, NT0909, AUSTRALIA

* Corresponding author: Centre for Marine Science and Technology’,

Department of Applied Physics, Curtin University of Technology, Bentley, WA 6102, AUSTRALIA

c.saIgado@cmst.curtin.edu.au

ABSTRACT

Sesarmid crabs are possibly one of the most important components of mangrove fauna because of their influence on
nutrient cycling and forest structure by feeding on litterfall. Little is known about the influence of clcctivity on the role of
crabs in mangrove forests, and how this is affected by the availability of litter items. This study investigated electivity of
three northern Australian sesarmid crabs ( Perisesarma semperi, Perisesanna darwinensis and Neosannatium meinerti)
from leaves of various conditions and from common species, as well as the effect of the availability of propagules
on feeding electivity. In almost every experiment, decayed and senescent leaves were selected over fresh leaves, and
typically decayed leaves were selected over senescent. Electivity for mangrove species, however, varied among crab
species and depended upon availability of litter type to select from. In experiments that included propagules as well
as leaves, leaves were selected over propagules. These results suggest that the sesarmid crab species included in this
study may have a greater role in nutrient cycling than in forest structuring because of their selectivity of leaves over
propagules.

Keywords: sesarmid, Sesarmidae, feeding behaviour, selectivity, mangrove leaves, consumption.

INTRODUCTION

Sesarmid crabs are one of the most common and
abundant faunal groups in mangrove forests (Golley et al.
1962; Jones 1984; Smith et al. 1991). Recent research
indicates that they play important roles in the ecology of
these ecosystems (Lee 1998) and may occupy a keystone
position in Australian mangrove forests (Smith et al.
1991). For example, crabs may affect forest structure by
attacking mangrove propagules (Smith 1987; McGuinness
1997), influence nutrient cycling by feeding on litterfall
(Robertson 1986), alter the properties of the soil by their
burrowing activities (Smith et al. 1991) and be involved
in competitive interactions with other species (Fratini et
al. 2000).

Studies on the feeding ecology of sesarmid crabs have
contributed to our understanding of the fate of mangrove
litter nutrients (Camilleri 1984, 1989). These studies
have shown that sesarmid crabs have an important role in
retaining nutrients within mangrove forests and reducing
export to nearby coastal systems (Lee 1997, 1998). Crabs
process a variety of food items, mainly dead leaves, into
smaller particles, and in this way make these nutrients
more readily available for other fauna to consume (e.g.
gastropods, crabs and other crustaceans). If sesarmid crabs
display selectivity for particular food items, such as leaves
and propagules from certain species of mangroves, this is

likely to affect the quantity, type and nutritional value of
mangrove litter that is recycled.

The feeding behaviour of some sesarmid crabs, in
particular those common to mangrove forests in north¬
eastern Australia and Kenya, has been studied (Camilleri
1989; Micheli 1993; Dahdouh-Guebas et al. 1997;
Kathiresan and Bingham 2001; Cannicci et al. 2007), but
the range of species and locations studied, is still limited.
Furthermore, previous studies did not test the effect of the
availability of litter on feeding electivity. The availability
of material may depend on factors such as mangrove
assemblage and season. In mangrove forests in tropical
Australia, for instance, most propagules drop during the
wet season (Ball and Pidsley 1988) and are, therefore, only
available at this time.

This study investigates feeding electivity of three
species of sesarmid crabs from common mangrove
species occurring in tropical mangrove forests in northern
Australia, and how electivity is affected by the seasonal
availability of propagules. The specific aim is to examine
electivity of mangrove leaves from three different species
and three conditions during wet and dry season conditions
(with and without the presence of propagules). Finally,
other variables which might affect consumption - crab
size (see Emmerson and McGwynne 1992) and sex (see
Olafsson et al. 2002) - were also investigated.

C. P. Salgado Kent and K. A. MeGuinness

MATERIALS AND METHODS

Crabs. The three most abundant sesarmid crabs in
the dominant mangrove assemblages in Darwin Harbour
(Salgado Kent 2004) were studied: Perisesarma semperi,
P. darwinensis, and Neosarmatium meinerti. Neosarmatium
meinerti is most abundant in mid to upper shore mixed
woodland and hinterland assemblages (Salgado Kent
2004) dominated by Ceriops australis and Avicennia
marina (naming of assemblages follows Brocklehurst and
Edmeades 1996). Perisesarma darwinensis is common
in tidal flat assemblages dominated by C. australis, but
P. semperi is found in tidal bank assemblages dominated
by A. marina and Rhizophora stylosa (Salgado Kent 2004;
pers. obs.). All crabs were collected from the forest at Jones
Creek, Darwin Harbour. Twenty individuals - ten of each
sex - of the two Perisesarma species were collected by
hand: half the crabs were larger, and half smaller, than
the average size (1.3 cm in carapace width). Average size
did not differ between species and was estimated from
106 crabs that were collected and measured prior to these
experiments. Neosarmatium meinerti crabs were captured
in tunneled pitfall traps (similar to the pitfall traps used
by Warren (1987)). Results for A. meinerti did not include
analyses on sex and size because only six individuals could
be captured, only one of which was female. All crabs were
placed in separate containers as soon as possible upon
return to the laboratory, to reduce stress and injury, in
particular among aggressive competing males.

Leaf preparation. The leaves and propagules used
in the experiments were taken from the three dominant
mangrove species in the four assemblages inhabited by
the crabs; A. marina, C. australis and R. stylosa. Leaf
conditions included fresh (green leaves), senescent (yellow
leaves) and decayed (brown leaves). Fresh and senescent
leaves were collected directly from trees. Fresh leaves can
frequently occur on the forest floor when storms, which are
common in the wet season, knock them down. Decayed
leaves were prepared by collecting senescent leaves from
trees and leaving these to decompose for fifteen days,
enclosed in 2 mm mesh bags tied to mangrove roots (as in
Robertson (1988)). All leaves w'ere stored at 4°C for 1 to
1.5 days, until the experiments began. Circular sections
of leaf, 2 cm in diameter, were used in experiments here
to reduce possible influences of leaf size on electivity (as
in Camilleri 1989). Propagules were collected from trees
and were also cut into similar sized pieces (to each other
and to the leaves).

Experiment preparation. All experiments were
conducted in a shaded, outdoor laboratory in which crabs
experienced a regular diurnal cycle, and conditions similar
to those prevailing in the field at that season. Crabs were
placed individually into clear, plastic containers, 14 cm
in diameter and 10 cm high. A circle 6 cm in diameter
in the centre of each container’s lid was cut out and this
allowed air to enter but prevented the crabs from escaping.

Containers without crabs were included in experiments to
control for weight changes due to leaching of dissolved
organic matter (DOM) and fungal and microbial activity.
Seawater was added to all containers to a level of 0.5 cm
and was changed daily (to ensure that lack of moisture
was not a factor affecting results, since the crabs inhabit
waterlogged environments). Most habitats where these
crabs were prevalent were saturated with water (with
the exception of N. meinerti). Crabs were starved for 24
hours before each experiment. During the experiments,
Perisesarma crabs were offered a choice of mangrove
material for a period of 12 hours. In each experiment,
the amount offered was one disc or piece for each type
x species (and each species x condition, for leaves) to be
included for testing. The amount of material offered was
great enough to allow for detection of significant patterns
in electivity, but small enough so that sufficient quantities
of material from discs and/or pieces remained to retain
information on patterns in electivity. For this reason N.
meinerti crabs were offered mangrove material for a
period of 18 hours, as these crabs took longer to consume
a significant amount of material. The material offered to
crabs was randomly placed within the containers. Each
batch of crabs was used in no more than two experiments to
ensure that their condition did not deteriorate significantly
and affect the results of the experiments.

Experiments. Two experiments were conducted
(Table 1). The first included leaves from all three mangrove
species and three leaf conditions, and the second included
the addition of propagules from all three mangrove
species. In mangrove forests of Darwin Harbour, Northern
Territory, the dominance of different mangroves species
differs among habitats where N. meinerti, P. semperi and
P. darwinensis are most abundant. However, all three
mangrove species overlap in distribution with the three crab
species to various extents (Brocklehurst and Edmeades
1996). Hence, in the natural environment, the chance of
encounter of leaves and propagules of these mangrove
species by the three species is realistic, and in most cases
relatively high.

To test for seasonal differences, experiments were
done under wet and dry season conditions (outdoor
laboratory experiments ensured that crabs experienced
seasonal changes in humidity and temperature). Dry season
experiments used only leaves but wet season experiments
used both propagules and leaves. A pilot study found
no significant effect of propagule dimension on feeding
electivity (Salgado Kent 2004). Each experiment described
below simultaneously examined feeding electivity of the
different species of crabs. ‘Dry Season’ experiments were
done simultaneously (and in the dry season), and ‘Wet
Season’ experiments were done simultaneously (in the
wet season).

Feeding selectivity of sesarmid crabs

Table 1. Material included in experiments testing electivity of sesarmid crabs for mangrove leaves and propagules. The mangrove species
were Am = Avicennia marina ; Ca = Ceriops australis; Rs = Rhizophora stylosa. The sesarmid crabs were Ps = Perisesarma semperi ; Sd =
Perisesarma darwinensis; Nm = Neosarmatium meinerti. Each experiment included six N. meinerti but twenty of each of the other species
(half male and half female; half small and half large). See the text for further details

Experiment 1: dry season

Electivity on common species

Experiment 2: wet season

Crab species

Leaf species

Fresh

Senesc

Senesc.

Senesc. Senesc.

Senesc.

Leaf condition

Decay

Propagule species

Experiment I. Electivity of leaves from three mangrove
species during dry season conditions. All crab species were
offered a choice of nine types of leaves, comprising all
combinations of the three species (C. australis, A. marina
and R. stylosa ) and three conditions (fresh, senescent, and
decayed).

Experiment 2. Electivity of leaves and propagules from
three mangrove species during wet season conditions.
Experiment 1 was repeated but with pieces of the
propagules of all three mangrove species also offered.

Processing measurements and calculations.
Consumption was measured in two ways, by weight
change and by area removed, and the results of these two
methods compared. The material offered had to be weighed
wet but the material remaining could be weighed wet or
dry. As dry weights were likely to be less variable, the
material remaining was dried at 60°C for two days and
then weighed. To calculate loss, the initial wet weights
were converted to dry weights. This was done by taking
wet and dry weights of twenty samples ol the leaves and
propagules of each species. These samples were then dried
at 60°C for two days and remeasured. Regression equations
predicting dry weight from wet were then derived and used
(all R2 > 0.8).

After the experiments, the area processed was calculated
by overlaying a clear plastic grid on top of each litter
item, counting the total number of 5 x 5 mm squares that
each leaf disc originally filled and the number of squares
consumed. The percent of area processed was converted
into dry weight processed by multiplying percent area by
the converted initial leaf dry weights.

The weight loss in control treatments (treatments with
no crabs) was subtracted from the weight loss of leaves and
propagules in treatments with crabs, to correct for weight
loss of propagules and leaves due to leaching of DOM and
fungal and microbial activity.

Statistical analyses. As the crabs were presented with
an array of choices simultaneously in each experiment,
the amounts of the different items consumed may not

have been independent (e.g. greater consumption of one
item is likely to result in reduced consumption of others).
Because of this, repeated measures analysis of variance
(ANOVA) was used for analysing all data in this study.
Assumptions for ANOVA were tested with Cochran’s
homogeneity of variances test and data were transformed
when appropriate. Mauchley’s Sphericity test was used
to check the assumptions required for repeated measures
ANOVA and, when these could not be met, the Greenhouse
Geisser correction was applied (Winer et al. 1991).

Several analyses were done: each included as many
factors as possible (chosen to test hypotheses a priori) so
as to limit the total number required. Data for P. semperi
and P. danvinensis in Experiments I and 2 could be
analysed together as the same numbers, sizes and sexes
of crabs were used and all were offered similar choices.
Analyses including crab size were based on absolute mean
consumption rates for the two groups (and was not weight
specific). Data for N. meinerti in Experiments 1 and 2 had
to be analysed separately as the choices offered to the
crabs differed. For all experiments, two sets of analyses
were done. One set compared the seasons but used only
the data for leaves (as propagules were not offered in the
dry season). The second set considered only the wet season
but included the data for propagules.

Factors in analyses varied, depending upon the design,
but included season (wet, dry), crab species ( P. semperi,
P. darwinensis), crab sex (male, female), size class of
crab (small, large), species of material (A. marina, R.
stylosa, C. australis) and type/condition of material
(fresh, senescent or decayed leaf; propagule). Some of the
resulting analyses were complex (i.e. Table 3). Inspection
of every higher order interaction in such analyses is likely
to be tedious and potentially unrewarding. Following
Mead (1988), our interpretation of such analyses focused
on sources of variation which were significant and which,
judged by the magnitude of the relevant mean square,
accounted for a substantial proportion of the variance.

C. P. Salgado Kent and K. A. McGuinness

0.15

0 13

Propagule

r 1 1 Fresh

0.04

- zz/> Senescent

■1 Decayed

0.02

0.00

A. marina

stylosa C. australis

Fig. 1. Consumption of material (g dry weight per 12 hours) from three mangrove species -A. marina, R. stylosa and C. australis - by N. meinerti
in electivity experiments (mean + SE). Note change in y-axis scale on graph for Experiment 2 (E2). For further details see Table 1.

RESULTS

Comparison of analyses based on leaf weight
processed and leaf area processed. Overall, results from
analyses based on weight and area were usually very
similar, although analyses of area consumed gave more
significant effects in the statistical analyses. This was
probably due to greater variability in estimates based on
weight (than on area). With measurements based on weight,
the initial observations, as they were of wet weights, were
likely to be more variable and the conversion of these
to dry weight, using the regression equations, probably
introduced additional errors. As there were relatively
few such differences, and they did not affect the overall
interpretation of the experiments, results in the rest of the
study are presented here from weights estimated from area
consumed.

Neosarmatium meinerti. Processing of leaves, in general,
depended upon the season, species and type/condition of
material when crabs were offered (see Table 2 for specific
factors affecting processing in each experiment; Figure

1) . The amount of decayed leaves processed was nearly
always greater than that of senescent or fresh leaves, with
the latter usually least preferred. Crabs processed more
R. stylosa than A. marina or C. australis, particularly in
the wet season when processing of some items increased
markedly (Figure 1: El v E2).

When propagules were offered with leaves (Experiments

2) , processing still depended upon the species and type/
condition of material (Table 2). Of the propagules, only
A. marina were processed and only in moderate amounts;
senescent and decayed leaves were processed in greater
quantities.

Perisesarma semperi. Patterns in processing of leaves
by P. semperi were complex, with numerous interactions
between the different factors in the analysis (Table 3).
Judged by the magnitude of the mean squares, the major
effects were of the species and type/condition of material,
and their interaction, and the interaction between crab size
and season. The overall pattern of results was very similar

to that seen with N. meinerti - greater processing of decayed
material than the other types, and of R. stylosa than of the
other species-although there was no increased processing
in the wet season (Fig. 2: El v E2). An interaction between
season and crab size (Table 3) occurred because large crabs
processed more leaf material in the dry season than small
crabs, but there was no difference between sizes in the
wet (means = 0.015 g/12 hours for large crabs in the dry
season, and 0.007 g/12 hours for small; 0.019 g/12 hours
for large crabs in the wet season, and 0.020 g/12 hours
for small). Interactions between size, season and other
factors (Table 3), however, indicate that the strength of this
pattern depends on the crab and mangrove species, and the
condition of the material. Interactions with crab sex were
also significant, although of less importance (Table 3).

When propagules were offered, the results were again
similar to those for N. meinerti : a moderate amount of
A. marina propagules was processed while other species

Table 2. Results of repeated measures analyses of variance of amount
of material processed (weights based on areas) by Neosarmatium
meinerti in all experiments using all common species (El, E2).
The table gives the df, MS and significance (* = P < 0.05;
*** = P < 0.001) for two analyses: the first used data for leaves only
but included two seasons (wet and dry); the second used data for
leaves and propagules but only for the wet season. Only the main
effects and significant interactions are shown. Effects with fractional
df have the Geisser-Greenhousc correction applied. See Table 1 for
further information on the design of the experiments.

Electivity for common species

Leaves only:

Leaves,

El, E2

propagules: E2

Factor

Season

1,9

0.032

Species (Sp) of
material

1.25, 18

0.076

2, 10

0.730*

Type/Condition of
material

2, 18

0.075*

3, 15

0.901***

Season x Sp

2, 18

0.013

Season x Type

2, 18

0.102*

Sp x Type

4, 36

0.025

6, 30

0.230

Season x Sp x Type

4, 36

0.066*

Feeding selectivity of sesarmid crabs

0.08

A. marina R. stylosa C. australis

Fig. 2. Consumption of material (g dry weight per 12 hours) from three
P. semperi in electivity experiments (mean + SE).

0.06

0.04

0.02

0.00

gggg Propagule
l l Fresh

A. marina R. stylosa C. australis

mangrove species - A. marina, R. stylosa and C. australis - by

were ignored (Fig. 2: E2). In Experiment 2, when materials
including propagules were offered, there was a substantial
effect of crab size (Table 3), but the interactions indicated
that the pattern was only observed for P. danvinensis (see
below).

Perisesarma danvinensis. Results for the consumption
of commonly available species by P. danvinensis were
broadly similar to those of the other two crab species (Table
3; Fig. 3) - the major effects were of the species and type/
condition of material and their interaction - but there were
differences in detail. While P. danvinensis still tended
to consume greater amounts of decayed leaves, greater

amounts of senescent and even fresh, leaves were taken
by this species (Fig. 3). In fact, in the dry season, roughly
equal amounts of all three types of C. australis leaves were
taken; and there was similar consumption of R. stydosa
decayed, and A. marina senescent and decayed, leaves
(Fig. 3: El). As with N. meinerti, there was a markedly
increased consumption of R. stylosa material in the wet
season (Fig. 3: E2). In contrast to the dry season, fresh and
senescent C. australis leaves were not taken at this time.

In Experiment 2, as noted above, when commonly
available materials, including propagules, were offered
there was a substantial effect of crab size (Table 3),

Table 3. Results of repeated measures analyses of variance of amount of material processed (weights based on areas) by Perisesarma semperi
and P. danvinensis in Experiments 1 and 2. Sec Table 1 for the design of the experiments and Table 2 for the format of the table.

Source

Leaves only:

El, E2

Leaves, propagules: E2

Season

1, 60

0.018

Crab species

1,60

0.055 *

1,28

0.198 *

Sex of crab

1,60

0.031

1,28

0.036

Size class of crab

1,60

0.009

1, 28

0.578 **

Species (Sp) of material

1.31, 120

0.337 ***

1.6, 56

1.416 ***

Type/Condition of material

1.46, 120

0.826 ***

2.25, 84

1.871 **

Season x Size

1, 60

0 152 ***

Sex x Sp

1.31, 120

0.032 *

1.6, 56

0.055

Sex x Type

1.46, 120

0.049 **

2.25, 84

0.049

Sp x Type

4, 240

0.140**

6, 168

1.004 ***

Season x Size x Sp

1.31, 120

0.054 **

Crab x Sex x Type

1.46, 120

0.036 *

2.25, 84

0.148 *

Season x Size x Type

1.46, 120

0.072 **

Crab x Size x Type

1.46, 120

0.005

2.25, 84

0.172 *

Sex x Size x Type

1.46, 120

0.024

2.25, 84

0.192*

Crab x Sp x Type

4, 240

0.045 ***

6, 168

0.071

Sex x Sp x Type

4, 240

0.023 *

6, 168

0.025

Season x Crab x Size x Type

1.46, 120

0.031 *

Season x Sex x Size x Type

1.46, 120

0.021 *

Crab x Sex x Sp x Type

4, 240

0.047 ***

6, 168

0.063

Season x Size x Sp x Type

4, 240

0.053 ***

Sex x Size x Sp x Type

4, 240

0.002

6, 168

0.166 **

Season x Crab x Sex x Size x Sp

1.31, 120

0.046 **

Season x Crab x Sex x Size x Type

1.46, 120

0.038 *

Season x Sex x Size x Sp x Type

4, 240

0.035 **

Season x Crab x Sex x Size x Sp x Type

4, 240

0.043 **

C. P. Salgado Kent and K. A. McGuinness

0.08 r

3 0.06 -

0.04

0.02

0.00

Jt lyJ

A. marina R. stylosa C. australis

0.14

0.12

0.10

0.08

0.06

0.04

0.02

0.00

Propagule
Fresh
ZZa Senescent
Decayed

A. marina R. stylosa C. australis

Fig. 3. Consumption of material (g dry weight per 12 hours) from three mangrove species - A. marina , R. stylosa and C. australis - by
P. darwinensis in electivity experiments (mean + SE). Note change in y-axis scale on graph for Experiment 2 (E2). For further details see
Table 1.

with large crabs processing more material than smaller
crabs (means = 0.013 g/12 hours, and 0.002 g/12 hours,
respectively).

DISCUSSION

A common trend in the present study was selectivity for
older material. When offered common species, N.meinerti
and P. semperi consumed more decayed R. stylosa leaves
than any of the other options (Table 4). Perisesarma
darwinensis displayed a similar pattern in the wet season,
although not in the dry. The second and third most consumed
materials were also usually decayed or senescent leaves,
although P. darwinensis electivity again differed in the dry
season. Other studies have documented similar electivity
for older material. In northeastern Australia, Camilleri
(1989) found Sesanna erythodactyla preferred aged leaves
to freshly fallen leaves and Micheli (1993) found Sesanna
messa preferred decayed leaves over senescent leaves. In
Kenya, Neosannatiwn smithii (= Sesanna smithii) preferred
old or decaying leaves over young leaves (Micheli 1991;
Table 5). Ashton (2002), however, found two Malaysian
species, Sesanna eumolpe and Sesanna onychophorum,
preferred fresh to senescent Avicennia officinalis leaves.

Studies have shown that decaying leaves have a lower
concentration of tannins than fresh leaves, so may be more

Table4. Summary ofelcctivity exhibited by the three speciesofsesarmids.
Am = A. marina ; Ca = C. australis; Rs = R. stylosa; f = fresh leaf;
s = senescent leaf; d = decayed leaf; p = propagule.

1st

2nd

3rd

Species

choice

Dry Season

N. meinerti

d-Rs

d-Am

d-Ca

P. semperi

d-Rs

d-Am

d-Ca,

s-Ca

P. darwinensis

d-Am

f-Ca

several

Wet Season

N. meinerti

d-Rs

s-Rs

d-Ca,

p-Am

P. semperi

d-Rs

s-Rs,

p-Am

P. darwinensis

d-Rs

s-Rs

d-Ca

easily digested and preferred for this reason (Giddins et
al. 1986; Neilson et al. 1986). A study by Micheli (1993),
testing for the effects of tannin, found no significant
correlation on the feeding preferences of N. smithii and S.
messa. These tests were, however, done with senescent,
rather than decaying, leaves (including R. stylosa,
C. australis and A. marina) and Robertson (1988) found
that the tannin content of these leaves decreased rapidly
over the first 14 days of decomposition. Thus, the range
of tannin concentrations in senescent leaves might not be
large enough to affect electivity.

Studies of sesarmid electivity for material from different
mangrove species, in contrast to material of different
ages, have given more variable results. Micheli (1993)
found that N. smithii preferred R. stylosa to A. marina ,
Bruguiera exaristata and C. australis; results similar to
P. semperi here. Camilleri (1989) found S. erythrodactyla
selected R. stylosa least, after A. marina and B. exaristata.
Ashton (2002), with S. eumolpe and S. onychophorum ,
found a preference for A. officinalis but only in fresh leaves.
With S. messa , however, Micheli (1993) found no significant
preference among the species tested and Dahdouh-Guebas
et al. (1997), studying N. meinerti in Kenya, also found
no preference, although only fresh material was offered.
Olafsson et al. (2002), however, also tested N. meinerti
from Kenya and obtained results similar to Camilleri
(1989) for 5. erythrodactyla. Greater electivity of A.
marina , as also exhibited on occasion by P. darwinensis
in the present study, can be explained by the particularly
low tannin and high nitrogen levels characteristic of this
species (Robertson 1988; Camilleri 1989; Michelli 1993).
Leaf nitrogen, in particular, is usually a reliable predictor of
herbivore preference in both laboratory and field situations
(Perez-Harguindeguy et al. 2003). Mature leaves from
A. marina in Darwin Harbour have been shown to have
significantly higher nitrogen concentrations than R. stylosa
and C. australis (Coupland 2002). Rhizophora stylosa
leaves, in contrast, have lower nitrogen concentrations and
higher percentage of tannin, than A. marina (Robertson
1988; Coupland 2002) and electivity for this species is
more difficult to explain.

Feeding selectivity of sesarmid crabs

Table 5. Summary of elcctivity/preferences for leaves of different species and conditions exhibited by sesarmid crabs in other studies.
Am = A. marina ; Ca = C. australis ; Ct = C. tagal; Bg Bruguiera gymnorltiza', Be = B. exaristata ; Rm = R. mucronata', Rs = R. stylosa ; Sa =
Sonneratia alba', f = fresh leaf; s = senescent leaf; d = decayed leaf; Nm = N. meinerti', Ns = N. smithi; Sm = 5. messa ; Se = S. erythrodactyla;
Cc = C. carnifex. Note: all species/conditions that were used in the trials are included below. *No pattern in electivity/preference.

Study

Field/lab

Species

1st choice

2nd choice

3rd choice 4th choice 5th choice

Steinke et al. 1993

Field

s-Bg

s-Am

f-Bg

f-Am

Giddins et al. 1986

Lab

d-Ct

s-Ct

f-Ct

Micheli 1993

Lab

s-Am, s-Rs, s-Ct, s-Be*

Lab

s-Rs

s-Am, s-Ct, s-Be

Lab

d-Am, d-Rs, d-Ct, d-Be

s-Am, s-Rs, s-Ct, s-Be

Field

Nm/ Sm

s-Ct

s-Rs, s-Re, s-Am

Camilleri 1989

Lab

d-Am

d-Bg

d-Rs

s-Am s-Bg; s-Rs

Lab

f-Am, d-Am

s-Am

Lab

d-Rs

s-Rs

Micheli etal. 1991

Lab

s-Bg

s-Sa

s-Rm

s-Ct s-Am

Lab

s-Bg. s-Sa, s-Rm, s-Ct, s-Am*

Camilleri (1989) found that crabs preferred thicker
R. stylosa leaves, so other leaf attributes may be influential
(such as moisture, fibre content and other chemical
constituents). Kennish and Williams (1997), in a study of the
tropical rocky shore crab Grapsus albolineatus, concluded
that algal morphology, through effects on feeding efficiency,
was more important than nutritional value or digestibility.
And Chavanich and Harris (2002) suggested that several
factors, including morphology and nutritional value,
probably influenced the feeding preferences of the subtidal
gastropod Lacuna vincta. Previous experience by the crabs
may also be important (Perez-Harguindeguy el al. 2003) and
explain some of the varying results for A. meinerti, which
has shown no particular pattern in electivity (Micheli et
al. 1991; Dahdouh-Guebas et at. 1997) and electivity for
(Olafsson et al. 2002) and against (Steinke et al. 1993; this
study) A. marina. It does not, however, appear to hold true
for crabs in the present study, since they did not usually,
when offered a range of common species, select species
from the assemblage in which they were most abundant
(in contrast to the results of Ashton (2002)).

Crab electivity within a restricted range of material,
representative of probable encounter rates, may differ from
electivity when all mangrove leaves and conditions are
offered in equal proportions. In a recent study on the gypsy
moth ( Lymatria dispar) for example, Raffa et al. (2002)
found that results could be affected by the combination
and arrangement of choices, and also by total consumption.
Perez-Harguindeguy et al. (2003) concluded that laboratory
preference experiments can predict relationships in
the field, but ecological factors such as variations in
accessibility and specialised plant-herbivore relationships
can cause differences. However, according to the model
they developed, predictions are likely to be reliable for
generalist herbivores and plants of high accessibility, both
conditions which apply here to various extents. Together,
these points suggest that strong, general trends, such as the
electivity for older material, are likely to apply in the field.

In contrast, more precise distinctions between species and
ages of material may be situation-specific. Further study
of weight-specific consumption rates, and experiments
offering different proportions of materials would shed light
on the different patterns observed here.

Given the importance of sesarmid crabs as propagule
predators (Smith 1987; McGuinness 1997; Lee 1998), the
limited consumption of propagules in the present study,
particularly by N. meinerti , is surprising. The Perisesarma
species may be too small to deal effectively with propagules
but N. meinerti is known to consume them in the field
(McGuinness 1997; Dahdouh-Guebas et al. 1997). On the
basis of these results, however, leaf material is preferred,
when it is available. Further, studies have found from 75%
(Steinke etal. 1993) to 90% (Dahdouh-Guebas et al. 1997)
of the material in N. meinerti stomachs to be leaf material,
although Skov and Hartnoll (2002) reported only 10%
of crabs feeding on leaves compared to 76% feeding on
mud. In part, the difference may result from the differing
digestibility of material: this would be consistent with
the results of Bouillon et al. (2002), whose stable isotope
studies indicated that sesarmid crabs fed on a wider range
of material than just mangrove leaves.

Effects of size and sex of crab were inconsistent,
although in accordance with previous results. Emmerson
and McGwynne (1992) reported a correlation between
size and consumption for N. meinerti in southern Africa.
Furthermore, Olafsson et al. (2002) found greater
consumption by female crabs, and they suggested this could
be due to either the difference in size between the sexes,
with females being smaller and having a higher potential
for energy loss, or to reproductive demands. The latter
explanation is perhaps more likely as size was controlled in
experiments here. Further, Micheli (1993) observed more
ovigerous crabs during the late dry season and this might
partially explain greater consumption at this time.

In conclusion, this study confirms the general pattern
in electivity of sesarmids for older material observed in

C. P. Salgado Kent and K. A. McGuinness

most other studies but also demonstrates that the range,
and types, of material offered can affect electivity. The
amount consumed depends upon the size of the crab, and
is likely to depend upon its sex and reproductive state.
Furthermore, the results suggest that sesarntid crabs
included in this study have a greater role in nutrient cycling
than in forest structuring because of their selectivity of
leaves over propagules. Hence, sesarmids in northern
Australia appear to have a distinct overall ecological role
than in other regions such as shown in some studies in new
world mangrove forests (Smith et al. 1991). Future studies
should attempt to confirm the findings here by; investigating
weight specific consumption rates, determining whether
‘feeding preferences’ differ from ‘feeding electivity’
(Underwood et al. 2004), and ultimately determining
the reasons underlying the high variability in sesarmid
feeding selectivity for leaves of different mangrove species,
observed in the majority of studies conducted thus far.

ACKNOWLEDGMENTS

This study was supported by a Large ARC grant (to K. A.
McGuinness). We thank other local mangrove researchers
for support and assistance. J. Warren provided helpful
advice on the design and use of pitfall traps. We offer
particular thanks to P. Davie for his valuable assistance
in the identification of Northern Territory sesarmid crabs.
Finally, the quality of the manuscript presented here has
been markedly improved by the input and insight of the
many reviewers, including J. Martin and K. Metcalfe.
The experiments presented in this study comply with the
current laws of the country in which the experiments were
performed.

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Accepted 8 October 2008

*•-

The Beagle. Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 33-54

Taxonomic review of Candalides absimilis (C. Felder, 1862) and C. margarita
(Semper, 1879) (Lepidoptera: Lycaenidae), with descriptions of two new

subspecies

MICHAEL F. BRABY

Biodiversity Conservation Division. Department of Natural Resources, Environment, the Arts and Sport, PO Box 496,

Palmerston, NT0831, AUSTRALIA

School of Botany and Zoology, The Australian National University, Canberra, ACT 0200, A USTRALIA

michael.braby@nt.gov.au

ABSTRACT

The taxonomic status of two members of the Candalides absimilis (C. Felder, 1862) species group from Australia is
reviewed. Two new subspecies, C. absimilis eastwoodi ssp. nov. from north-eastern Queensland and C. absimilis edwardsi
ssp. nov. from south-eastern Australia, are described, diagnosed and compared with related taxa. C. gilberti Waterhouse,

1903 from north-western and central northern Australia is shown to comprise a geographical race of C. margarita
(Semper, 1879) based on comparative evidence of the genitalia, adult and immature stage morphology, life history
and adult behaviour, and accordingly is treated as a subspecies of that taxon. Information on the distribution, habitat,
conservation status and biology are summarised for each of the three taxa, with additional notes provided on the larval
food plant, behaviour and life cycle of C. absimilis edwardsi ssp. nov, which specialises as larvae on the flush growth
of Bracfiychitan populneus (Schott and Endl.) R.Br. (Sterculiaceae) growing in eucalypt woodland. Females of both
C. absimilis edwardsi ssp. nov. and C. margarita gilberti are remarkable in that they comprise the only taxa within the
C. absimilis species group from Australia in which the white patches on the fore and hind wing are replaced with blue. A
striking association between habitat (broad vegetation type) and adult female phenotype in the C. absimilis species group
is noted, with highly contrasting ‘black and white forms’ occurring in closed forest and less striking ‘blue forms’ in open
forest/woodland, and this relationship is found to extend more generally within the Australian Lycaenidae. A general
hypothesis, that ambient light properties among divergent habitats or vegetation types (i.e. different light environments)
is a potent selective force shaping sex-limited phenotype, is proposed for these butterflies.

Keywords: ambient light environment, Australian monsoon tropics, butterfly, Candalidini, Candalides, Cape York Peninsula, Iron
Range, East Gippsland, sexual dimorphism.

INTRODUCTION

The tribe Candalidini of lycaenid butterflies includes
about 33 species referred to two genera, Candalides Hiibner
and Nesolycaena Waterhouse and Turner, both of which
are endemic to the Australian Zoogeographic Region (Tite
1963; Edwards and Kerr 1978; Braby 1996, 2000; Braby
and Douglas 2004). Braby (2000) and Braby and Douglas
(2004), building on the foundational work of Tite (1963)
and Edwards and Kerr (1978), recognised three species
groups within Candalides, viz : the C. absimilis (C. Felder,
1862) species group, the C. erinus (Fabricius, 1775) species
group, and the C. xanthospilos (Hiibner, [1817]) species
group. Phylogenetic relationships among the three species
groups of Candalides and Nesolycaena are unknown at
present, but each taxonomic unit is assumed to comprise
a monophyletic group. By far the largest species group is
the C. absimilis species group, containing 21 described
species (Parsons 1998; Braby 2000; Tennent 2006). The
group is characterised by having a comparatively long labial
palpus, especially in the female; the androconia! scales in

the male fore wing being concentrated into a diffuse trident¬
shaped patch along the basal half of veins M v CuA, and
CuA„ as well being distributed on the adjacent veins; and
the underside pattern of the wings possessing a series of
small dark markings and wavy lines but usually without
terminal spots, although some species have one or more
black tomal spots on the hind wing. In males, the termen of
the hind wing is often produced towards the tomus, and the
upperside ground colour is lustrous blue; while in females,
the upperside ground colour of both wings is frequently
black with a prominent white central patch. Members of this
species group are distributed predominantly in rainforest
habitats of mainland New Guinea and its adjacent islands
and the coastal areas of eastern Australia. One species also
extends across the monsoon tropics of northern Australia,
while another, C. cvprotus (Olliff, 1886), occurs in
heathland habitats in the coastal, semi-arid and arid areas
of southern, central and south-western Australia (Braby
2000; Grand 2006).

In the past few decades, six species in the Candalides
absimilis species group have been recognised from the

M. F. Braby

Australian mainland (Common and Waterhouse 1972,
1981; Edwards 1996; Braby 2000; Edwards et al. 2001).
One of these, C. absimilis, shows complex geographical
variation, with divergent populations at the extreme
northern and southern ends of the range, and three
distinct forms were recognised by Braby (2000). Another
taxon, C. margarita gilberti Waterhouse, 1903 stat. rev.
from the Kimberley and Top End of north-western and
central northern Australia, has hitherto been treated as
a distinct species since its description, but Braby (2000)
postulated that it may be conspecific with C. margarita
(Semper. 1879), comprising a geographical subspecies
closely related to the nominate subspecies from eastern
Australia. The purpose of this paper is to clarify and
revise the taxonomic status of the geographical forms
of C. absimilis and the taxonomic status of C. margarita
gilberti. Information is also summarised on the distribution,
habitat, conservation status, and biology of each species.
Attention is drawn to a striking association between adult
female wing colour pattern and light characteristics among
divergent habitats (broad vegetation types) in Candalides ,
and this relationship is found to extend more broadly within
the Australian Lycaenidae.

METHODS

The male and female genitalia of Candalides absimilis
were dissected and examined from six populations spread
across the species’ latitudinal range, viz: Cape York
Peninsula (Iron Range, Mt White Coen); Wet Tropics
(Shiptons Flat, Townsville); central Queensland (Mackay,
Eungella); south-eastern Queensland (Eudlo); north-eastern
New South Wales (Broken Head); and eastern Victoria
(Mitchell River). The male and female genitalia of C.
margarita were examined from four populations distributed
across the geographical range of the species within
Australia, viz: Top End of the Northern Territory (Darwin,
Adelaide River); Gulf Country of western Queensland
(Musselbrook, near Lawn Hill National Park); Wet Tropics
(Paluma Range west of Townsville); and north-eastern New
South Wales (Grafton).

The size of the white central patch in female Candalides
absimilis was analysed quantitatively across the species
geographical (latitudinal) range. Specimens (N= 173) from
seven populations spread along the eastern coast, viz: (1)
Cape York Peninsula (Iron Range); (2) Wet Tropics (Caims-
Townsville); (3) central Queensland (Mackay-Eungclla);
(4) south-eastern Queensland (Brisbane); (5) central New
South Wales (Sydney); (6) south-eastern coastal New
South Wales (Central Tilba-Bega); and (7) eastern Victoria
(East Gippsland), were measured to assess the extent of
geographical variation. Thirty specimens were sampled
from each population, except for Cape York Peninsula and
central Queensland for which only eight and 15 specimens
were available respectively. The shape of the white patch on

both wings is either an ellipse or approximately circular; the
area of the patch was therefore calculated using the formula
Tt^r,, where r, = radius 1, and r,=radius 2. For the fore wing,
r, was measured along the cubitus and vein M 3 , while r was
measured perpendicular to r p from the mid costa to tornus;
for the hind wing, r, was measured along vein M„ while r,
was measured perpendicular to r,, from the apex to tornus.
Radial measurements were made with vernier callipers to
a precision of 0.1 mm. Fore wing length, from the base to
the apex of the wing, was measured for each specimen as
a proxy for body size.

The following abbreviations refer to repositories where
material has been examined:

AMS: Australian Museum, Sydney

ANIC: Australian National Insect Collection,
Canberra

BMNH; Natural History Museum, London (formerly
British Museum of Natural History)

MCZ: Museum of Comparative Zoology, Harvard

University, USA

MFBC: private collection of Michael Braby

MTQ: Museum of Tropical Queensland,

Townsville

NMV: Museum Victoria, Melbourne (formerly

National Museum of Victoria)

NTM: Museum and Art Gallery of the Northern

Territory, Darwin (formerly Northern Territory
Museum)

SAM: South Australian Museum, Adelaide

Label data of all material are given as depicted on the
specimen, with quotation marks used to designate each
separate label for specimens with two or more labels. The
symbol “|” is used to denote text on the reverse side of a
label.

TAXONOMY

Candalides absimilis eastwoodi ssp. nov.

(Figs 7-10, 26)

Type material. HOLOTYPE - c? labelled “MT.
WHITE, COEN, 7 JAN 1994, R. EASTWOOD” (ANIC).
PARATYPES- 14cfc?, 399. QUEENSLAND; 19 labelled
“CLAUDIE R., 29 12 13”, “LEP-9841” (NMV); 19
labelled similarly but with date “18 2 1914”, registration
number"LEP-9837” (NMV); 19 labelled similarly but with
date “5 4 14”, registration number “LEP-9840” (NMV);
2c? labelled “Claudie. R. Nth. Q., 5/16-V-61, J. Macquecn”
(ANIC); 1 cf labelled “Mount White, Coen, Qld, I3°58’S
143° 11 ’E, 29 Apr 1989, G. and A. Daniels”, “G. Daniels
Collection” (AMS); 4c? labelled “Mt White, Coen, Qld,
13°58’S 143°] l’E, 7 Jan 1994 447m, G. and A. Daniels”,
“G. Daniels Collection” (AMS); 3cf labelled “MT. WHITE,
COEN, 7 JAN 1994, R. EASTWOOD” (ANIC); 4c?
labelled similarly but with date “12 JAN 1994” (ANIC).

Taxonomic review of Candalides

Figs 1-25. Type material of Candalides absimilis and C. margarita: 1-3, Holochila absimilis C. Felder, 1862 lectotype male from Ash Island,
NSW (BMNH), showing upperside, underside and specimen labels; 4-6, C. persimilis Waterhouse, 1942 lectotype male from Mackay, Qld
(AMS), showing upperside, underside and specimen labels; 7-8, C. absimilis eastwoodi ssp. nov. holotype male from Mount White, Coen,
Qld (ANIC), showing upperside and underside; 9-10, C. absimilis eastwoodi ssp. nov. paratype female from Iron Range. Qld (NMV),
showing upperside and underside; 11-12. C. absimilis edwardsi ssp. nov. paratype male from Mitchell River National Park, Vic. (NMV),
showing upperside and underside; 13-14, C. absimilis edwardsi ssp. nov. holotype female from Suggan Buggan. Alpine National Park,
Vic. (ANIC), showing upperside and underside; 15-16, 23, Holochila margarita Semper, 1879 lectotype male from Bowen, Qld (BMNH),
showing upperside, underside and specimen labels; 17-18, C. margarita margarita female from Bluewater State Forest west of Townsville,
Qld (MFBC), showing upperside and underside; 19-20, 24, C. gilberti Waterhouse, 1903 lectotype male from Darwin, NT (AMS), showing
upperside, underside and specimen labels; 21-22,25, C. gilberti paralectotype female from Darwin, NT (AMS), showing upperside, underside
and specimen labels. Scale bar = 20 mm.

M. F. Braby

Other material examined. 23cfcT, 699-
QUEENSLAND: 1U labelled “1 mile NE Mt. Lamond,
Iron Range. Qld., 21 Dec. 1971”, “D.K. McAlpine, G.A.
Holloway, D.P. Sands” (AMS); 19 labelled “Claudie R,
2 miles S Mt. Lamond, Iron Range, Qld., 13 Jan. 1972”,
“D.K. McAlpine & G.A. Holloway”, “Australian Museum
K231664” (AMS); 19 labelled “ Candalides consimilis
toza? 9, Claudie Rv area. N.Q.. 4 January 1974, Shane F
McEvey.”, “Genitalia Slide M345” (ANIC); lcf labelled
“Iron Range, Cape York. Qld, 13.xi. 1991, S.J. Johnson”
(MTQ); lef labelled “IRON RANGE, N. Qld, 25 Nov. 1995,
D J FERGUSON” (ANIC); 2cf, 1 9 labelled “Iron Range,
Cape York Pen. Qld., 5-11 .xii. 1995, S.J. Johnson” (MTQ);
19 labelled “Wcnlock River crossing, Coen-Iron Range
road, Cape York Pen., N. Qld., 10 Sep. 1974, M.S. & B.J.
Moulds”, “M.S. MOULDS COLLECTION”, “Australian
Museum K231665” (AMS); 19 labelled “Rocky River (nth
of Silver Plains), Qld, 13°48.415’S, 143 o 28.20UE, 27 Jul
2005, R.P. Field”, “LEP-480S8” (NMV); lcMabelled “Mt
White, Coen, N. Qld, 5 Nov. 1979, M.S. & B.J. Moulds”,
“M.S. MOULDS COLLECTION” (AMS); 1 cf labelled
“Mount White, Coen, Qld, 13°58’S 143°1 HE, 29 Apr 1989,
G. and A. Daniels”, “G. Daniels Collection” (AMS); 1 cf
labelled “Mt. White, Coen. Qld., 19,i. 1993, S.J. Johnson”
(MTQ); 1 cf labeled “Mt White, Coen, Qld, 10 Jul 1993,
R.P. Field”, “LEP-44854” (NMV); 1 cf labelled similarly
but with date “11 Jul 1993” and registration number “LEP-
44855” (NMV); 1 cf, 19 labelled “Mt White, Coen, Qld,
13°58’S 143°1 l’E, 7 Jan 1994 447m, G. and A. Daniels”,
“G. Daniels Collection” (AMS); 1 cf labelled similarly
but with date and collector “12 Jan 1994, G. Daniels R.
Eastwood” (AMS); 2cT labelled “MT. WHITE, COEN, 7
JAN 1994, R. EASTWOOD” (MCZ); 3cf labelled “Mt.
White, Coen. QLD., 27.iv. 1997, S.J. JOHNSON” (MTQ);
3cf labelled similarly but with date “9.v. 1998” (MTQ); 1 cf
labelled “Mt White, Coen, Qld, I3°58.035’S, 143°11.538’E,
4 Jul 2005, R.P. Field”, “LEP-48086” (NMV); 1 cf labelled
similarly but with registration number “LEP-48087”
(NMV); 1 cf labelled similarly but with date “5 Jul 2005”
and registration number “LEP-48085” (NMV).

Adult description. Male. Fore wing length 16.4-19.2
mm. Upperside of wings lilac-blue, with costa, termen and
outer half of dorsum narrowly edged black, radial, median,
cubital and anal veins black towards wing margin; fore wing
with a broad distinct median patch of androconial scales,
patch more prominent at anterior end of cell and base of
veins M : and M, and their origin with the discocellulars,
at base of veins M,, CuA, and CuA, and their origin with
the cubital vein (Cu), and along vein IA+2A; hind wing
with a conspicuous black tomal spot between ends of veins
CuA, and 1A+2A, and a smaller indistinct black terminal
spot between ends of veins CuA, and CuA,. Underside
of wings silvery white, with a series of small dark brown
markings and a narrow dark brown termen; hind wing with
a conspicuous dark brown tomal spot between ends of veins
CuA, and 1A+2A, and sometimes a series of four smaller

indistinct dark brown terminal spots between ends of veins
M, and CuA,. Female. Fore wing length 17.2-19.8 mm.
Upperside of wings black enclosing a large white central
patch, base suffused with purplish blue scales; fore wing
with purplish blue scales extending along posterior margin
of white central patch above vein 1A+2A and sometimes
along anterior margin of patch in cell; hind wing with
scattered purplish blue scales bordering posterior and outer
margins of white central patch. Underside of wings with
ground colour and markings similar to male, but markings
usually more distinct.

Variation. Males show minor variation in the extent
of the tomal spot on the hind wing, and in the shape of the
hind wing in which the termen and tomus are sometimes
rounded. In females, the extent of the white central patch
on the upperside of the hind wing varies.

Comparison. Candalides absimilis eastwoodi ssp. nov.
males are distinct and can be readily distinguished from
males of C. absimilis absimilis (Figs 1 -6) and C. absimilis
edwardsi ssp. nov. (Figs 11-12) by differences in the shape
of the fore wing in which the termen is straighter and less
convex, and the apex more sharply pointed; the shape of
the hind wing in which the tornus is generally produced to
a point; the extent of the underside markings, which are less
conspicuous and only faintly visible, especially the spots
and wavy lines on the hind wing; the upperside ground
colour, which is paler purplish blue; the median patch of sex
scales on the fore wing, which is more clearly visible; and
the extent of the black/dark brown tomal spot on the hind
wing, which is generally more prominent on the upperside
and always present on the underside. Candalides absimilis
absimilis males from the northern end of their range (i.e.
Wet Tropics - between Cooktown and Townsville) are
similar to C. absimilis easbvoodi ssp. nov. in that the tornus
of the hind wing is often produced and the tomal spot is
usually present (though usually not as well developed),
but otherwise more closely resemble the lectotype male of
C. absimilis absimilis from Ash Island, New South Wales
(Figs 1, 2), in that the termen of the fore wing is more
rounded, the underside markings are more distinct, and
the upperside ground colour is brighter/deeper blue with
the patch of sex scales less clearly visible.

Candalides absimilis eastwoodi ssp. nov. females
differ from C. absimilis edwardsi ssp. nov. females by
possessing a large, conspicuous white central patch on the
upperside of each wing. The white patch on the fore wing is
similar in size to that of C. absimilis absimilis populations
from south of the Wet Tropics (i.e. Mackay-Sydney), but
apparently smaller than those from the Wet Tropics (i.e.
Cooktown-Townsville). The white patch on the hind wing
of C. absimilis eastwoodi ssp. nov., however, is substantially
larger than populations of C. absimilis absimilis from south
of the Wet Tropics, but significantly smaller than those
from the Wet Tropics (see quantitative analysis below
under Remarks).

Taxonomic review of Candalides

Candalides absimilis eastwoodi ssp. nov. has previously
been confused with C. consimilis Waterhouse, 1942 (Dunn
et al. 1994; Dunn 1995) to which it closely resembles.
On Cape York Peninsula, C. consimilis is known only
from Iron Range and has been separated as the northern
subspecies C. consimilis toza Kerr, 1967. Dunn et al.
(1994) initially placed material from Mount White near
Coen with C. consimilis toza, which is known only from a
few male specimens (Kerr 1967), but later considered them
to belong to an undescribed subspecies of C. consimilis
closely allied to C. consimilis toza (Dunn 1995). However,
the valva of the male genitalia (Fig. 26) and sinus vaginalis
of the female genitalia (not illustrated) clearly indicate
that the population from Mount White belongs with C.
absimilis and not with C. consimilis. Also, males of C.
absimilis eastwoodi ssp. nov. are distinguished from those
of C. consimilis toza by the absence of a conspicuous
series of black terminal spots on the uppersidc of the fore
and hind wings, particularly between the ends of veins
M, and CuA,.

Remarks. Waterhouse (1942) described Candalides
persimilis from Mackay, Qld, and placed material from the
Wet Tropics (Cairns, Kuranda) and coastal north-eastern
New South Wales (Byron Bay) with that species. The taxon
was differentiated on the basis of the paler lilac uppersidc
ground colour of the male with a more conspicuous patch
of sex-scales, a large white central patch on the fore wing
of the female, the presence of a distinct black tornal spot on
the underside of the hind wing, and minor differences in the
male genitalia in which the apices of the valvae are smaller
and truncate. Titc (1963) and McCubbin (1971) questioned
the validity of C. persimilis Waterhouse, 1942, but Tindale
(1965) recognised the taxon and indicated a wide sympatric
distribution with C. absimilis along the eastern coast, from
Cairns, Queensland, to Narara, New South Wales. Tindale
crudely illustrated and compared the male genitalia,
although his Figure 4 does not match that of Waterhouse’s
(1942) Figure 1 d for the same structural character. Common
and Waterhouse (1972, p. 415) did not recognise the species,
noting that “A close study of C. persimilis Waterhouse
suggests that it is simply a form of C. absimilis." The
species group name persimilis was subsequently treated as
a junior synonym of C. absimilis by Edwards (1996) and
Edwards el al. (2001). Examination of the lectotype male
(Figs 4-6) and paralectotypes of C. persimilis in the AMS,
including the male genitalia, indicate that this synonymy
is justified. The minor differences in the genitalia alluded
to by Waterhouse (1942) and Tindale (1965) are part of the
overall geographical variation observed in C. absimilis.
The male genitalia from three widely dispersed populations
(Cape York Peninsula, south-eastern Queensland, eastern
Victoria) are illustrated in Figures 26-28. The genitalia show
minor geographical variation, particularly in their overall
size and in the shape of the valvae. In the northernmost
population (C. absimilis eastwoodi ssp. nov.), the genitalia
are consistently smaller, and the bifurcated apices of the

valvae are smaller, less pronounced and rotated inwardly
so that they lie perpendicular to the length of the valva
(Fig. 26). The genitalia from the Wet Tropics (C. absimilis
absimilis) appear transitional with those from Cape York
Peninsula (C. absimilis eastwoodi ssp. nov.) and Mackay-
Eungella in central Queensland (C. absimilis absimilis) in
having relatively small apices that are partially rotated.

Quantitative analysis of the size of the white central
patch of Candalides absimilis females (,V= 140) revealed
that patch size, when present, was positively correlated
with body size, and this relationship was more tightly
correlated in the fore wing (r = 0.755, d.f 138, P < 0.001)
than in the hind wing (r= 0.560, d.f 138, P< 0.001) (Fig.
33). After controlling for the effects of body size, patch
size was found to exhibit clinal variation, being negatively
correlated with latitude (Fig 34). Analysis of variance of
mean patch size, measured for both fore and hind wing
from the seven populations sampled across the species’
geographical range, revealed that the relationship was
highly significant for each (fore wing: F= 7.348, d.f. 5, P
= 0.0422; hind wing: F= 11.990, df 5 ,P = 0.0180). Closer
inspection of the relationship, however, suggests a steep
cline in this character for the hind wing but not the fore
wing between Caims-Townsville and Mackay-Eungella,
Qld, negligible change between Mackay, Queensland, and
Sydney, New South Wales, followed by another steep cline
approaching a step for both wings south of Sydney, New
South Wales, to East Gippsland, Victoria. (Fig. 34). Patch
size reached its maximum extent in the Wet Tropics, and
then appeared to diminish in size further north on Cape
York Peninsula, although the sample size for the latter
population (n = 8 for C. absimilis eastwoodi ssp. nov.) was
too small for comparison with the Wet Tropics. Although
C. absimilis absimilis females from the Wet Tropics have
the largest white patches on both wings, this population
is better treated as a local form rather than as a distinct
subspecies until more C. absimilis eastwoodi ssp. nov.
females become available for comparison. C. absimilis
absimilis females from Mackay more closely resembled
populations from Brisbane and Sydney than those from
Caims-Townsville with respect to patch size (see also
comments by Tite 1963).

Etymology. The taxon is named in honour of Dr
Rodney Eastwood who has made a substantial contribution
to the scientific study of Australian butterflies, especially
the natural history and ecology of ant-lycaenid associations
and the phylogcography of Lycaenidae.

Distribution. Candalides absimilis eastwoodi ssp.
nov. is restricted to Cape York Peninsula, from Iron Range
(McEvey 1977) south along the Mcllwraith Range to the
Rocky River north of Silver Plains (R.P. Field) and Mt
White near Coen, Queensland (Braby 2000) (Fig. 35). It
is known from six sites (i.c. point localities > 1 km apart)
representing four discrete locations (i.e. areas > 10 km
apart). The subspecies is probably endemic to the mid¬
peninsula rainforest block (Iron Range-Mcllwraith Range)

M. F. Braby

1 mm

Figs. 26-32. Male genitalia of Candalides absimilis and C. Margarita , showing ventral view, with aedeagus removed: 26. C. absimilis eastwoodi
ssp. nov. from Iron Range, Old; 27, C. absimilis absimilis from Eudlo, Old: 28. C. absimilis edwardsi ssp. nov. from Mitchell River, Vic.; 29,
C. margarita gilberti from Darwin, NT; 30, C. margarita gilberti from Musselbrook, Gulf of Carpentaria, Qld; 31, C. margarita margarita
from the Paluma Range west of Townsville, Qld; 32, C. margarita margarita from Grafton, NSW. Scale bar = 1 mm.

Taxonomic review of Candalides

Body size (mm)

Fig. 33. Relationship between body size (measured as fore wing
length) and size of white patch for each wing in Candalides absimilis
females: A, fore wing; B, hind wing. Correlations are significant for
both wings (P < 0.001).

of Cape York Peninsula, which is geographically separated
from the Wet Tropics where the nominate subspecies
C. absimilis absimilis reaches its northernmost limit at
Cooktown, Queensland.

Despite reports to the contrary, Candalides absimilis
eastwoodi ssp. nov. has not been recorded further north
than the tip of Cape York Peninsula and the Torres Strait
islands. These reports were found, on closer inspection, to
be erroneous. A male labelled “Prince of Wales Is., 9 June
1908, H. Elgner.”, “LEP-9890”, “Seems to belong with
Candalides absimilis Feld., det K.L. Dunn 1989” (NMV)

is considered to have been mislabelled and excluded from
the material examined. Dunn and Dunn (1991) considered
the record to be reliable and included Prince of Wales
Island from southern Torres Strait islands in their review
of the distribution of the species. A detailed chronology
of Elgner’s geographical movements in the Torres Strait
islands and Cape York, based on insect specimens that he
collected (Moulds 1977; Dunn 2007), indicate that for
the period 28 April-19 July 1908, Elgner was stationed
on Prince of Wales Island, so he would have had access
to the island in June 1908. However, Elgner’s specimen
of C. absimilis does not agree with the characteristics
of C. absimilis eastwoodi ssp. nov. It lacks the straight
termen and pointed apex of the fore wing, paler upperside
ground colour, and indistinct markings on the underside; it
more closely resembles C. absimilis absimilis from south¬
eastern Queensland and New South Wales. Moreover,
examination of the genitalia indicates that it conforms
with typical C. absimilis absimilis (Fig. 27) and not to C.
absimilis eastwoodi ssp. nov. (Fig. 26). Dunn (2007) noted
a high level of accuracy in Elgner’s label data; therefore,
it is possible the specimen was mislabelled by subsequent
curators. Two male specimens of C. margarita margarita
in AMS labelled “Horn Is. Q„ 25.6.43” and “Cape Yk.
Q, 25.6.43” were found to have been misidentified
as C. absimilis in the drawers of the main reference
collection.

Biology. The early stages and larval food plants of
Candalides absimilis have not been recorded. Adults have
been collected in September, from November to February
and in April, May and July, with most specimens collected
during the ‘build-up’ and early wet season. Available
data on adult phenology suggests the subspecies is on
the wing throughout the year. Near Coen, males have
been collected hilltopping at the summit of Mount White
(R.G. Eastwood, personal communication; D.A. Lane,
personal communication), a behaviour that appears to
be largely absent in the two other subspecies (except
for the population of C. absimilis absimilis in the Wet
Tropics, which has been recorded hilltopping at Mt Stuart
Townsville, M.F. Braby unpublished data).

Conservation status. Candalides absimilis eastwoodi
ssp. nov. has a narrow geographical range, with an estimated
extent of occurrence of4,500 km 2 . Available data suggests
the subspecies is restricted to tropical forest of the mid¬
peninsula rainforest block of Cape York Peninsula (i.e. Iron
Range-Mcllwraith Range), most of which is now protected
in the Iron Range and Mcllwraith National Parks. There
is no information on population trends, fragmentation of
populations, or immediate threatening processes impacting
on the known sites, and the ecology of the subspecies is
very poorly known. Although the range of the taxon is
suspected to be relatively circumscribed, there is inadequate
information to make a direct assessment of the risk of threat.
Its conservation status should therefore be regarded as Data
Deficient (DD) according to 1UCN criteria.

M. F. Braby

Fig. 34. Relationship between size of white patch divided by body size and latitude for both fore and hind wing in Candalides ahsimilis females
sampled from seven populations across the species’ geographical range. Points are mean values with error bars representing one standard
deviation. Sample sizes are shown above graph. Regression equations for the significant relationship between mean patch size and latitude
are as follows: y = 1.984 - 0.0388.V for fore wing (F 7.348, d.f 5 ,P = 0.0422), and y 1.733 - 0.0409.V for hind wing (F 11.990, d.f 5,
P = 0.0180). Accompanying map of eastern Australia shows distribution of C. absimilis (modified from Braby 2000), with examples from
each of the seven geographical areas, as follows: A, Cape York Peninsula, Qld (Iron Range-Mcllwraith Range); B, Wet Tropics, Qld (Cairns-
Townsville); C, central Queensland (Mackay-Eungclla): I), south-eastern Queensland (Brisbane): E, central New South Wales (Sydney);
F, south-eastern coastal New South Wales (Central Tilba-Bcga); and G, eastern Victoria (East Gippsland).

Candalides ahsimilis edwardsi ssp. nov.

(Figs 11-14, 28)

Type material. HOLOTYPE - 9 labelled “36°56’S,
148°21’E; 600m, 7 km N Suggan Buggan (by rd), Alpine
NP, VIC., emg. 6 JAN. 2003, M.F. Braby & L.J. Aitchison”,
“Reared from larva on Brachychiton populneus, coll. 8-9
DEC. 2002, pupated 25 DEC. 2002“ (AN1C). PARATYPES
- 15CTCT, 1499. NEW SOUTH WALES: 19 labelled
“36°45’S, 148°25’E; 300m, 18 km N Willis (by road),
Kosciuszko NP, NSW; emg. 13 JAN. 2003, M.F. Braby
& LJ Aitchison”, “Reared from larva on Brachychiton
populneus , coll. 10 DEC. 2002, pupated 2 JAN. 2003”
(AMS). VICTORIA: 19 labelled “36°56’S, 148°21 ’E;
600m, 7 km N Suggan Buggan (by rd), Alpine NP, VIC.,
emg. 4 JAN. 2003, M.F. Braby & L.J. Aitchison”, “Reared
from larva on Brachychiton populneus, coll. 8-9 DEC. 2002,
pupated 23 DEC. 2002” (NMV); lef labelled similarly but
with dates “emg. 5 JAN. 2003”, “pupated 22 DEC. 2002”
(AMS); 19 labelled similarly but with dates “emg. 12
JAN. 2003”, “pupated 31 DEC. 2002” (AMS); 1 Cf labelled
similarly but with dates “emg. 15 FEB. 2003”, “pupated 29
DEC. 2002” (NMV); lef labelled similarly but with dates
“emg. 17 OCT. 2003”, “pupated 30 DEC. 2002” (ANIC);

IcT labelled “37°05'S, 148°24’E; 200m, McKillops Bridge,
Snowy River NP, VIC., 3 JAN. 2004, M.F. Braby” (ANIC);
1 cf labelled “37°05’S, 148°24’E, McKillops Bridge, Snowy
RiverNP, VIC., emg. 23 JAN. 2004, M.F. Braby”, “Reared
from larva on Brachychiton populneus, coll. 3 JAN. 2004”
(NMV); I cf labelled similarly but with date “emg. 27 JAN.
2004” (AMS); 1 cf labelled similarly but with date “emg.
28 JAN. 2004” (AMS); 19 labelled similarly in NMV; 1 cf.
I 9 labelled but with date “emg. 29 JAN. 2004” (ANIC);
1 cf labelled similarly in NMV; 19 labelled similarly in
AMS; IcT labelled “37°29’S, 148°09’E, Buchan Caves
Reserve, VIC; 100m, emg. 10 JAN. 2004, M.F. Braby”,
“Reared from larva on Brachychiton populneus, coll. 19-21
DEC. 2003” (ANIC); 1 cf labelled similarly but with date
“emg. 12 JAN. 2004" (AMS); 19 labelled similarly in
NMV; 1 cf labelled similarly but with date “emg. 15 JAN.
2004” (NMV); 19 labelled similarly but with date “emg.
16 JAN. 2004” (ANIC); 19 labelled “37°42’S, I47°21’E,
Den of Nargun, Mitchell River NP, VIC.., emg. 7 JAN.
2003, M.F. Braby & L.J. Aitchison”, “Reared from egg on
Brachychiton populneus, coll. 6-7 DEC. 2002, pupated 24
DEC. 2002” (ANIC); 1 cf labelled similarly but with dates
“emg. 9 JAN. 2003”, “pupated 28 DEC. 2002” (NMV);

Taxonomic review of Candalides

distribution of Candalides absimilis eastwoodi ssp. nov. (■).

1 cT labelled similarly but with date “emg. 10 JAN. 2003”
(ANIC); 19 labelled similarly but with dates “emg.
12 JAN. 2003”, “pupated 30 DEC. 2002” (AMS); 19
labelled similarly but with date “pupated 31 DEC. 2002”
(NMV); 19 labelled “37°42’S, 147 0 21’E, DenofNargun,
Mitchell River NP, VIC., 23 DEC. 2003, M.F. Braby &
L.J. Aitchison” (ANIC); 19 labelled similarly in AMS;
19 labelled “37°42’S, 147°21’E, Den of Nargun, Mitchell
River NP, VIC., emg. 15 JAN. 2004, M.F. Braby”, “Reared
from larva on Brachychiton populneus , coll. 21-23 DEC.
2003” (NMV); lef labelled similarly but with date “emg.
19 JAN. 2004” (AMS).

Other material examined. 61 cfcf, 329 9- NEW SOUTH
WALES: lef labelled“36°45’S, 148°25’E; 300m, 18kmN
Willis (by road), Kosciuszko NP, NSW; 10 DEC. 2002, M.F
.Braby & L.J. Aitchison” (ANIC); 1 cf labelled similarly in
NMV; lef labelled similarly in AMS; lef labelled “36°45’S,
148°25’E; 300m, 18 km N. Willis, Kosciuszko NP, NSW;
emg. 3 MAR. 2003, M.F. Braby”, “Reared from larva on
Brachychiton populneus, coll. 10 DEC. 2002, pupated 2
JAN. 2003” (MFBC). VICTORIA: lef labelled “36°56’S,
148°21’E; 600m, 7 km N Suggan Buggan (by rd), Alpine
NP, VIC., emg. 4 JAN. 2003, M.F. Braby & L.J. Aitchison”,
“Reared from larva on Brachychiton populneus, coll. 8-9
DEC. 2002, pupated 22 DEC. 2002” (MFBC); 19 labelled

similarly but with date “pupated 23 DEC. 2002” (MFBC);
19 labelled “37°05’S, 148°24’E, McKillops Bridge,
Snowy River NP, VIC., emg. 29 JAN. 2004, M.F. Braby”,
“Reared from larva on Brachychiton populneus, coll. 3
JAN. 2004,” (MFBC); lef labelled similarly but with date
“emg. 30 JAN. 2004” (MFBC); 19 labelled similarly but
with dates “emg. 30 AUG. 2004, pupated late JAN. 2004”
(MFBC); 19 labelled similarly but with date “emg. 10 SEP.
2004” (NMV); lef labelled “Buchan Caves, Buchan, Vic,
37°30.16’S, 148°9.51’E, 11 Jan 2003, R.P. Field”, “LEP-
46133” (NMV); 1 cf labelled similarly but with registration
number “LEP-46134” (NMV); lef labelled similarly but
with registration number “LEP-46135” (NMV); 19 labelled
“Buchan Caves, Buchan, Vic, 37°29.63’S, 148°10.08’E, 13
Dec 2003, R.P. Field”, “LEP-46197” (NMV); 19 labelled
similarly but with date “17 Jan 2004” and registration
number “LEP-46201” (NMV); 19 labelled similarly but
with date “ern 20 Feb 2004” and registration number “LEP-
46202” (NMV); 19 labelled similarly but with date “cm 27
Feb 2004” and registration number “LEP-46203” (NMV);
lef labelled similarly but with registration number “LEP-
46204” (NMV); 1 cf labelled similarly but with date “em 22
Sept 2004” and registration number “LEP-46789” (NMV);

1 cf labelled similarly but with registration number “LEP-
46790” (NMV); 1 cf labelled similarly but with date “em 29
Sept 2004” and registration number “LEP-46785” (NMV);

1 cf labelled similarly but with registration number “LEP-
46786” (NMV); 1 cf labelled similarly but with date “em 5
Jan 2005” and registration number “LEP-46799” (NMV):

1 cf labelled similarly but with registration number “LEP-
46800” (NMV); 1 cf labelled similarly but with date “em 25
Jan 2005” and registration number “LEP-46903” (NMV);
lef labelled similarly but with registration number “LEP-
46904” (NMV); 1 cf labelled similarly but with registration
number “LEP-46905” (NMV); lef labelled similarly but
with registration number“LEP-46906” (NMV); 19 labelled
similarly but with date “em 26 Jan 2005” and registration
number “LEP-46901” (NMV); 1 9 labelled similarly but
with registration number“LEP-46902” (NMV); 1 cf labelled
similarly but with date “em 27 Jan 2005” and registration
number “LEP-46897” (NMV); lef labelled similarly but
with registration number“LEP-46898” (NMV); 1 cf labelled
similarly but with registration number “LEP-46899”
(NMV); 19 labelled similarly but with registration number
“LEP-46900” (NMV); 19 labelled similarly but with date
“em 28 Jan 2005” and registration number “LEP-47218”
(NMV); 19 labelled similarly but with registration number
“LEP-47219” (NMV); 19 labelled similarly but with date
“em 29 Jan 2005” and registration number “LEP-47220”
(NMV); 1 9 labelled similarly but with date “em 30 Jan
2005” and registration number “LEP-47221” (NMV); 19
labelled similarly but with registration number “LEP-47222”
(NMV); 19 labelled "37°29’S, 148°09’E, Buchan Caves
Reserve, VIC; 100m, emg. 11 JAN. 2004, M.F. Braby”,
“Reared from larva on Brachychiton populneus, coll. 19-
21 DEC. 2003” (MFBC); 19 labelled similarly but with

M. F. Braby

date “emg. 12 JAN. 2004” (MFBC); lcT labelled similarly
but with date “emg. 17 JAN. 2004” (MFBC); lcf labelled
similarly but with dates “emg. 17 FEB. 2004. pupated DEC.
2003” (MFBC); 1 cf labelled similarly but with dates “emg.
10 JUL. 2004, pupated JAN. 2004” (MFBC); Id labelled
similarly but with dates “emg. 12 SEP. 2004, pupated late
DEC. 2003” (MFBC); Id labelled “VIC Den ofNargun,
40 km NE Stratford, 37°42’S 147°21'E, 7 December
1977, Shane F. McEvey”, “S.F. McEvey B1 OOOd” (AMS);
Id labelled similarly but with registration number “S.F.
McEvey B100Id” (AMS); Id labelled similarly but with
registration number “S.F. McEvey B1002d” (AMS); Id
labelled “VIC The Amphitheatre, Billy Goat Bend, Mitchell
River, 37°40’S 147°22’E, 8.xii. 1977 S.F. McEvey”, “S.F.
McEvey B1008d” (AMS); Id labelled similarly but with
registration number “S.F. McEvey B1009d” (AMS);
Id labelled similarly but with registration number “S.F.
McEvey BlOlOd” (AMS); Id labelled similarly but with
registration numbers “S.F. McEvey B1011 d”, “Australian
Museum K233023” (AMS); 19 labelled similarly but with
registration numbers “S.F. McEvey B10069”, “Australian
Museum K233021” (AMS); 19 labelled “VIC 1km upriver
from The Amphitheatre on Billy Goat Bend, Mitchell
River, 37°39’S 147 0 21’E, 9.xii.l977 S.F. McEvey” “S.F.
McEvey B10209”, “Australian Museum K233024”
(AMS); 1 d labelled similarly but with registration number
“S.F. McEvey B102Id” (AMS); Id, 29 labelled “DEN
OF NARGUN, Mitchell River, Vic., 17 Dec. 1989, K.L.
Dunn, J.M. Dunn” (ANIC), d with additional label “AN1C
genitalia slide No. 3183 d” and genitalia preparation “LYC
3183, Candalides absimilis (Feld.), AUST. ENT. INS.
COLL. Slide E.D. Edwards 1990”, 9 with additional label
“Holotype, Candalides pseudogoodingi”; Id, 19 labelled
“VIC., Den ofNargun, Mitchell River N.P., 17 Dec. 1989,
K.L. & J.M. Dunn”, “Candalides absimilis, det KL Dunn
1989”, “Reference: DUNN 1990, Victorian Entomologist
20: 49-53” (ANIC); 19 labelled similarly in SAM; Id
labelled similarly but with registration number “LEP-
9732” (NMV); Id labelled similarly but with registration
number “LEP-9733” (NMV); 1 9 labelled similarly but
with registration number“LEP-9754” (NMV); 1 9 labelled
similarly but with registration number “LEP-9734” (NMV);
Id labelled “DEN OF NARGUN. V., MITCHELL R.
N.P.,21 FEB. 1991, D.F. CROSBY”(ANIC); Id labelled
“37°42’S, 147°21’E, Den ofNargun, Mitchell River NP,
VIC., emg. 1 JAN. 2003, M.F. Braby & L.J. Aitchison",
“Reared from larva on Brachychiton populneus, coll. 6-7
DEC. 2002, pupated 19 DEC. 2002” (MFBC); 19 labelled
similarly but with dates “emg. 5 JAN. 2003, pupated 22
DEC. 2002” (MFBC); 1 d labelled similarly but with date
“pupated 23 DEC. 2002” (MFBC); Id labelled similarly
but with dates “emg. 7 JAN. 2003, pupated 24 DEC. 2002”
(AMS); Id labelled similarly but with dates “emg. 8
JAN. 2003, pupated 25 DEC. 2002” (MFBC); 19 labelled
similarly but with dates “emg. 10 JAN. 2003, pupated 30
DEC.2002”(MFBC); Id, 19 labelled“37°42’S, 147°2UE,

Den ofNargun, Mitchell River NP, VIC., 2 JAN. 2004,
M.F. Braby” (ANIC); 2d, 19 labelled similarly in NMV;
4d labelled similarly in AMS; 3d labelled similarly in
MFBC; Id labelled “37°42’S, I47°21’E, Den ofNargun,
Mitchell River NP, VIC., emg. 12 JAN. 2004, M.F. Braby”,
“Reared from larva on Brachychiton populneus , coll. 21 -23
DEC. 2003” (NMV); Id, 19 labelled similarly but with
date “emg. 13 JAN. 2004” (MFBC); 2d labelled similarly
but with date “emg. 14 JAN. 2004” (MFBC); Id labelled
similarly but with date “emg. 19 JAN. 2004 (ANIC); Id
labelled similarly but with date “emg. 23 JAN. 2004”
(MFBC); 19 labelled similarly but with dates “emg. 18
SEP. 2004. pupated JAN. 2004” (MFBC); Id labelled
similarly but with dates “emg. 23 SEP. 2004, pupated 11
JAN. 2004” (MFBC).

Adult description. Male. Fore wing length 15.5 - 16.6
mm. Upperside of wings purplish blue, with costa and
termen narrowly edged black, radial, median, cubital and
anal veins black towards wing margin; fore wing with a
broad indistinct median patch of androconial scales, patch
more prominent at anterior end of cell and base of veins
M, and M, and their origin with the discocellulars, at base
of veins M,, CuA, and CuA, and their origin with the
cubital vein (Cu), and along vein 1A+2A; hind wing with
a small black tornal spot between ends of veins CuA, and
1A+2A. Underside of wings silvery white, with a series
of dark brown markings and a narrow dark brown termen;
hind wing usually with an obscure dark brown tornal spot
between ends of veins CuA, and 1A+2A, and a series of
smaller indistinct dark brown terminal spots between ends
of veins Rs and CuA,.

Female. Fore wing length 15.3- 16.9 mm. Upperside of
wings with costa, termen and outer half of dorsum broadly
black enclosing a large purplish blue central patch which
extends from base and inner half of dorsum to postmedian
area; fore wing usually with a small postmedian patch or
suffusion of pale whitish blue scales distal to discocellulars
between veins M, and CuA! or CuA,. Underside of wings
with ground colour and markings similar to male.

Variation. The brightness of the upperside purplish blue
ground colour varies in males. In females, the purplish blue
central patch on the upperside occasionally extends narrowly
as a suffusion of purplish blue scales to the costa between
veins R, and R, and to the subterminal area between veins
CuA, and 1A+2A on the fore wing, and more broadly to
the termen between veins M, and 1A+2A on the hind wing.
The upperside of the hind wing very occasionally possesses
a small subapical patch or suffusion of pale whitish blue
scales between veins Rs and M, or M r

Comparison. Candalides absimilis edwardsi ssp. nov.
females are distinguished from females of C. absimilis
absimilis and C. absimilis eastwoodi ssp. nov. by
pronounced differences in the extent of the white central
patch on the upperside. In C. absimilis edwardsi ssp. nov.,
the white patch on the fore wing is absent, being replaced
with a small postmedian patch of pale whitish blue scales

Taxonomic review of Candahdes

between veins M, and CuA,; occasionally this patch is
restricted to a narrow postmedian streak between veins
M, and M, distal to the discal cell. The white patch is also
absent on the hind wing, but occasionally there is a narrow
subapical whitish blue streak between veins Rs and M r
Some C. absimilis absimilis females from south-eastern
coastal New South Wales (Central Tilba-Bega) approach
C. absimilis edwardsi ssp. nov. in having much reduced
or no white central or subapical patch on the upperside of
the hind wing and an extensive basal suffusion of purplish
blue scales, but these specimens have a more conspicuous
white central patch on the fore wing that is approximately
circular in shape and which extends from veins M, to CuAj
or CuA,.

Candalides absimilis edwardsi ssp. nov. males do not
differ from those of nominate C. absimilis absimilis in
external morphology. The male genitalia (Fig. 28), however,
are slightly larger, with the apical projections of the valvae
more pronounced.

Candalides absimilis edwardsi ssp. nov. females are
similar to those of C. eonsimilis goodingi (Tindale, 1965)
from eastern Victoria, but the latter are readily distinguished
by the more pointed apex of the fore wing, less rounded
tomus of the hind wing, presence of a small but conspicuous
white central patch on the upperside of the fore wing which
extends below vein M r and the well-defined basal spots
on the underside of the hind wing. In addition, the central
iridescent area in C. eonsimilis goodingi is a brighter,
richer blue, whereas in C. absimilis edwardsi ssp. nov. it is
purplish blue. The two species also differ significantly in
their genitalia (see Waterhouse 1942; Tite 1963).

Remarks. Candalides absimilis edwardsi ssp. nov.
was previously regarded as a distinct local form that was
believed to be restricted to the Mitchell River National
Park of East Ciippsland (Dunn 1990; Dunn and Dunn
1991; Dunn et al. 1994). Dunn and Dunn (1991, p. 391)
concluded that in relation to the white central patch of
females “This Victorian population represents the opposite
extreme of the continuum ... at the subspecific level this
population would be considered a poorly differentiated
taxon”. However, quantitative analysis of this character
over the broad geographical range of the species (Fig. 34)
indicates negligible variation between Mackay-Eungella,
Queensland, and Sydney, New South Wales, but a steep cline
approaching a step between Sydney, New South Wales, and
East Gippsland, Victoria. C. absimilis edwardsi ssp. nov.
lies at the southernmost limit of the species’ geographical
range and represents the extreme end of variation in the
extent of the white patch. Its disjunct distribution, together
with a broader geographical range than hitherto believed,
and the striking female phenotypic divergence in which
the reduced white patch comprises a consistent and
diagnosable character state that lies at the end of a step or
steep latitudinal cline, indicate that the known populations
warrant formal recognition at the subspecific level.

Examination of label data of museum material of
Candalides absimilis edwardsi ssp. nov. revealed several
inconsistencies. The original Dunn material comprised
10 specimens (4cf, 69) collected from Den of Nargun on
17 December 1989. Most specimens have an additional
label "Candalides absimilis, det KL Dunn 1989”;
however, one female in the ANIC is labelled in Dunn’s
original handwriting “Holotype” and a manuscript name
of Dunn’s is associated with it. Because this name was
never formally published, neither the specimen nor the
name carry any nomenclatural validity. A male sent to
the ANIC for taxonomic identification (see Dunn 1990)
has its accompanying genitalia slide labelled “LYC 3183,
Candalides absimilis (Feld.), Den of Nargun, Mitchell
River, Vic., 17 Dec. 1989, K.L. Dunn & J.M. Dunn, AUST.
ENT. INS. COLL. Slide E.D. Edwards 1990”, indicating
that correct taxonomic diagnosis of the species was not
made until 1990.

Etymology. The taxon is named in honour of Mr Ted
Edwards in recognition of his outstanding contribution to the
study ofAustralian Lepidoptera, curation and development
of the ANIC, and continuous support, generous help and
mentoring to others, including the author, over many
decades.

Fig. 36. Map of south-eastern Australia showing geographical
distribution of Candalides absimilis edwardsi ssp. nov. (■) and
C. absimilis absimilis (A). Stars (★) indicate occurrence of the
natural larval food plant, Brachychiton populneus, of both subspecies
in the region (data based on records registered in Australia’s Virtual
Herbarium http://www.cpbr.gov.au/cgi-bin/avh.cgi).

M. F. Braby

Distribution. Candalides absimilis edwardsi ssp. nov.
is known from a limited region in south-eastern Australia
(Fig. 36). It has been recorded from three disjunct areas
in south-eastern New South Wales inland of the Great
Escarpment and East Gippsland of eastern Victoria at
altitudes between 100-600 m. These three areas include:
(1) the Upper Snowy River area on the New South Wales-
Victoria border (M.F. Braby); (2) Buchan Caves Reserve
(New et cd. 2007); and (3) Mitchell River National Park
(Dunn 1990). In the Upper Snowy River area, it has been
recorded from three locations, viz: the southern sections of
Kosciuszko National Park, New South Wales, close to the
Victorian border; at Suggan Buggan and Willis in Alpine
National Park, Victoria; and at McKillops Bridge in Snowy
River National Park, Victoria. At Buchan Caves Reserve, it
is known only from one site. In the Mitchell River National
Park, it is known from three sites along the Mitchell River:

1 km upstream from The Amphitheatre on Billy Goat Bend,
The Amphitheatre on Billy Goat Bend, and Den ofNargun.
Thus, C. absimilis edwardsi ssp. nov. is known from a total
of eight sites representing five locations.

Candalides absimilis edwardsi ssp. nov. is geographically
separated from C. absimilis absimilis , which reaches its
southernmost limit in the coastal/subcoastal areas of south¬
eastern New South Wales (Fig. 36). C. absimilis absimilis
was previously knowrn as far south as Tilba Tilba, New
South Wales (Braby 1998), but surveys by the author
have detected the species further south near Cobargo and
several sites at Bcga. The two subspecies are separated
by a minimum distance of 120 km in south-eastern New
South Wales, but are divided by the Great Escarpment
and montane plateau of the Monaro Plains. C. absimilis
absimilis occurs east of the escarpment in moist coastal
eucalypt open-forest, whereas C. absimilis edwardsi ssp.
nov. occurs w'est of the escarpment in the dry box woodlands
in the rain-shadow area of the Upper Snowy River south
of the Great Dividing Range. The natural larval food plant
of both subspecies throughout this region is Brachychiton
populneus (Schott and Endl.) R.Br. (Braby 1998, and
unpublished data), although at 13 km west of Cobargo,
G. Guy (personal communication 2007) has also recorded
C. absimilis absimilis breeding on several planted (non-
indigenous) rainforest trees, viz: Macadamia integrifolia
Maiden and Betche, Harpullia pendula Planchon ex
F. Muell., and Stenocarpus sinuatas Endl. The latter two
species have not previously been recorded as larval food
plants for C. absimilis , although near Brisbane females were
observed ovipositing on new shoots of Harpullia pendula
(Braby 2000).

Comparison of the spatial distribution of
Brachychiton populneus in south-eastern Australia with
that of C. absimilis shows a close geographical relationship
(Fig. 36). The larval food plant is apparently absent from
the Monaro Plains (between Bredbo and Bombala, New
South Wales) west of the Great Escarpment, suggesting that
the two butterfly subspecies are indeed isolated. Searches

for C. absimilis edwardsi ssp. nov. by the author in the
relatively dry woodland areas inland of the Great Dividing
Range at Mount Majura and the Murrumbidgee River
corridor, Australian Capital Territory, and the Wee Jasper
district. New South Wales, where Brachychiton populneus
grows abundantly failed to detect the butterfly. Presumably
C. absimilis edwardsi ssp. nov. is also absent from the upper
reaches of the Murray River inland of the Divide where
the food plant likewise is present. Dunn and Dunn (1991)
recorded C. absimilis absimilis from the Australian Capital
Territory, based on a single female (lodged in the ANIC)
collected from the Brindabella Range, but the phenotype of
this specimen does not match specimens from south-eastern
coastal New South Wales, suggesting the specimen is
mislabelled. C. absimilis absimilis in south-eastern coastal
New South Wales is otherwise known only from the coastal
areas east of the Great Escarpment.

Four specimens of Candalides absimilis recorded from
three localities in the outer Melbourne region (Healesville,
Dandcnong Ranges, Mordialloc) registered in the AMS and
NMVare considered to be erroneous and accordingly their
data excluded from the material examined. These specimens
are labelled as follows: 1 cf “Dandcnong Ranges, Victoria |
3765”, "KL16576”, “G.A. Waterhouse Collection” (AMS);
19 “Dandcnong Ranges, Victoria | C. French, 3766”,
“KL 16576”, “KL22003”, “G.A. Waterhouse Collection”
(AMS); 19 “HEALESVILLE, J.A.K.” (NMV); 19
“MORDIALLOC, J.A.K.” (NMV). In addition, there is a
female specimen in NMV reputedly from Victoria labelled
“J.A. Kershaw, Collection, PURCH JUNE 1941”. In all
cases, the females resemble typical C. absimilis absimilis
in that they have a prominent white central patch on the
upperside of each wing and do agree with populations of C.
absimilis edwardsi ssp. nov. from eastern Victoria in which
the white patches are absent. Waterhouse (1942), Tindale
( 1 965) and Dunn (1990) referred to this material in part and
regarded the specimens as authentic, but Dunn and Dunn
(1991) seriously questioned their validity and regarded the
label data as unreliable. Dunn (1990) concluded that the
specimens resulted from multiple accidental introductions
to Melbourne but which failed to establish a resident
breeding population, but he did not identify the mechanism
or source of the introductions. However, the inadequate data
on the labels and limited number of specimens suggest the
material is probably mislabelled.

Crosby (1951) recorded ''Candalides absimilis' from
eastern Victoria based on observations at Gypsy Point,
Victoria; however, these records refer to C. consimilis
goodingi, the species of which was not widely recognised
at that time and which had previously been confused
with C. absimilis (Waterhouse 1942). Crosby (1951) also
indicated that material of‘C. absimilis' had been collected
from the northern reaches of the Macalister River north of
Heyfield (by V. Smith), but this material was subsequently
described by Tindale (1965) as Holochila goodingi, with
Macalister River as the type locality. McEvey (1979)

Taxonomic review of Ccindalides

Figs 37-57. Habitat and life histories of Candalides absimilis and C. margarita: 37, eucalypt woodland habitat of C. absimilis edwardsi ssp. nov.
on the slopes ofthc Mitchell River, Vic., showing larval food plant Brachychiton populneits in deciduous form; 38, eucalypt woodland habitat
of C. absimilis edwardsi ssp. nov. at McKillops Bridge, Snowy River, Vic., showing larval food plant Brachychitonpopulnens in foreground
with new flush growth; 39. eucalypt woodland habitat of C. absimilis edwardsi ssp. nov. at Suggan Buggan. Vic; arrow indicates larval food
plant Brachychiton populneits with new flush growth; 40, open woodland habitat of C. absimilis edwardsi ssp. nov. at Buchan Caves, Vic.,
showing larval food plant Brachychiton populneus with mature foliage; 41. Brachychiton populneits new leaf growth and flowers; 42. egg
of C. absimilis edwardsi ssp, nov.; 43, egg of C. margarita gilberti ; 44, egg of C. margarita margarita ; 45, early instar larva of C. absimilis
edwardsi ssp. nov.; 46-47, final instar larva of C. absimilis edwardsi ssp. nov. showing dorsal and lateral views and colour forms; 48-49. pupa
of C. absimilis edwardsi ssp. nov. showing dorsal and lateral views; 50-51, final instar larva of C. margarita gilberti showing dorsal and lateral
views; 52-53, pupa of C. margarita gilberti showing dorsal and lateral views; 54-55, final instar larva of C. margarita maigarita showing
dorsal and lateral views; 56-57, pupa of C. maigarita margarita showing dorsal and lateral views.

M. F. Braby

recorded C. consimilis goodingi from the Mitchell River
during surveys in 1977-78; however, careful examination
of this material (13 specimens in AMS) indicated a mixed
series comprising two species - three specimens (399) of
C. consimilis goodingi and 10 specimens (8cfcF, 299) of
C. absimilis edwardsi ssp. nov. McEvcy surveyed three sites
(Den of Nargun, The Amphitheatre on Billy Goat Bend, 1
km upstream from The Amphitheatre on Billy Goat Bend)
over a three day period (7-9 December 1977). C. absimilis
edwardsi ssp. nov. was recorded from each of these sites,
but C. consimilis goodingi was collected only at The
Amphitheatre, indicating that the two taxa occur together
but presumably with limited overlap. The Mitchell River
is the only known location where these two species are
sympatric in Victoria. McEvey’s material from the Mitchell
River thus appears to represent the first genuine collection
of C. absimilis from Victoria.

Habitat. In the Upper Snowy River area (Figs 38, 39),
the breeding population occurs in dry eucalypt woodland,
chiefly dominated by Eucalyptus albens Benth. (White
Box) with some Callitris glaucophylla Joy Thomps. and
L.A.S.Johnson (White Cypress Pine) and Eucalyptus
melliodora A.Cunn. ex Schauer (Yellow Box) in the valley
of the Snowy River, including banks of the river itself as
well as gentle slopes several hundred metres above the river.
In this habitat, the larva! food plant is sparsely distributed
over a relatively wide area, often occurring as single isolated
trees, compared with those growing on the slopes of the
Mitchell River (Fig. 37). The area lies in a pronounced
rainfall shadow in which the mean annual rainfall (c. 800
mm) is much lower than the coastal areas of New South
Wales of similar latitude further east.

At Buchan Caves Reserve (Fig. 40), the breeding
population occurs in open woodland on limestone outcrops
on exposed slopes with a north or west facing aspect, or
along the valley floor of Fairy Creek, where the larval
food plant grows. The larval food plants vary greatly in
size and age, from very old large trees to young saplings
only a few metres high and, within the Reserve itself, occur
in relatively low density (a total of only 16 plants were
located). A few scattered trees (mostly ornamental garden
and street trees) also occur in residential areas of Buchan
close to the Reserve where the butterfly also breeds.

At Mitchell River National Park (Fig. 37), the breeding
population occurs in woodland dominated by Brachychiton
populneus and Eucalyptus melliodora, with a sparse
understorey of shrubs and small trees of Acacia implexa
Benth. (Lightwood) and A. mearnsii De Wild. (Black
Wattle), and grasses. The habitat occurs as a narrow strip on
both sides of the Mitchell River in a limited zone where the
larval food plant grows abundantly on steep rocky slopes
between the top of escarpment and the bottom of the gorge/
river valley (i.e. Mitchell River and its tributaries, such as
Woolshed Creek). This habitat type is patchy in distribution
because the food plants generally grow on rocky slopes
with a predominantly a north to west facing aspect where

conditions are drier and more exposed. As a result, the
larval food plants are more concentrated and less widely
dispersed than those occurring in the Upper Snowy River
area. Warm temperate rainforest dominated by Tristaniopsis
laurina (Sm.) Peter G.Wilson and J.T.Waterh., Acmena
smithii (Poir.) Merr. and L.M.Perry, Elaeocarpus reticulatus
Sm., Pittosporum undulation Vent, and Myrsinehowittiana
(Mcz) Jackcs prevails along the gorges; however, the larval
food plant does not grow, and the butterfly does not breed,
in this habitat.

Biology. Larval food plant. Brachychiton populneus
(Schott and Endl.) R.Br. (Sterculiaceae) (Kurrajong). The
larval food plant typically grows as a tree up to 10 m in
height, with a dense, dark green crown of foliage. Long¬
term observations made on the phenology of Brachychiton
populneus in south-eastern Australia by the author indicated
that the trees flowered and regenerated their foliage over
a short interval during summer, mainly in December and
January, although some trees produced new flushes of
growth as late as February. Although foliage regeneration
was very seasonal and occurred rapidly among individual
trees, not all trees regenerated synchronously, and there was
a succession of new annual leaf growth during the summer
period. A proportion of trees were also deciduous (Fig.
37) - some plants dropped all of their leaves during late
spring and early summer before regeneration, especially in
dry years or those growing in the more exposed situations.
Other trees produced new growth before shedding most of
their older mature leaves, and thus remained vegetated.

Behaviour. Eggs (Fig. 42) were laid singly or sometimes
in pairs, usually on the stem immediately below a terminal
branchlet of new developing leaf buds, or on the new soft
shoots and stipules of the young developing leaves. Many
eggs were laid on individual trees, but females oviposited
only on trees producing flushes of new growth or, for those
trees that had shed all of their older leaves, in the early
phases of regeneration; they generally did not oviposit on
trees in which the leaves had reached an advanced phase
of regeneration. Females were observed to oviposit on the
same tree over a minimum period of 10 days. Individual
trees probably remain suitable foroviposition for about two
to three weeks due to the rapid growth of new foliage.

Early instar larvae (Fig. 45) fed singly on the new soft
leaves (Fig. 41) at night, resting by day on the underside
of the young leaves where they avoided direct sunlight.
Usually only one larva occurred on a single terminal cluster
of foliage, but occasionally up to four larvae were found.
The later instar larvae, when not feeding, typically rested
lower down on the petiole of a leaf where they remained
well concealed, often in the fork between the petiole and
the branchlet. Between instars, the larvae settled on the
leaf petiole or further down on the thicker stems of the
branchlet to moult. In the late instars (Figs 46, 47), larvae
varied considerably in colour, from uniformly green to
almost entirely reddish with some green at the posterior
end; others were intermediate between these two phenotypic

Taxonomic review of Candalides

extremes, being green with a red mid-dorsal line and a red
lateral line and a slight pinkish tinge at the anterior end.
Variation in larval colour was associated with the colour
of juvenile foliage, which was either bright red or bright
green (Fig. 41). Larvae were not observed attended by
ants, despite the abundance of Crematogaster sp. ants on
many trees.

Larvae, when reared in captivity, showed a preference
to pupate in well-concealed, dark situations, such as inside
rolled bark. The pupa was attached, to a silken platform
spun over the substrate, by anal hooks and a silken central
girdle. Pupae (Figs 48, 49) were noted to stridulate when
stimulated (e.g. when gently stroked or sprayed with water).
In the field, pupae were not found on the foliage, flowers or
trunk and branches of the larval food plant, and it is likely
that the larvae leave the food plant to pupate amongst dry
leaf litter (dead leaves, bark etc) or under rocks, stones or
inside rock crevices at the base of the tree.

In captivity, adults always emerged during the morning,
well after sunrise, but usually before midday. After
emergence, adults quickly expanded their wings and were
ready for flight within 30-60 mins. In the field, males were
noted to fly rapidly throughout much of the day in the mid-
to upper canopy, usually between 2-8 m above the ground,
around the outer foliage of larger trees of the larval food
plant. They typically patrolled around a tree and then quickly
flew to the next; they were also observed to fly around the
outer foliage of Pittospomm undulatum, the leaves of which
arc superficially similar to Brachychitonpopulneus. Males
rarely alighted to settle, especially when many individuals
were present, but once settled, they usually perched for a
few seconds, or sometimes for up to about 1 min, high up
on horizontal leaves of the outer foliage of the larval food
plant. When fewer individuals were present, males perched
for longer periods on the upper foliage, especially during
the late afternoon, to establish territories. In these situations,
males would leave their perch briefly to patrol around the
crown of the tree before settling again, usually in a different
place on the tree. When settled, the wings remained closed
or half opened at an angle of about 90° towards the sun.
The patrolling and perching behaviour of males suggest that
newly emerged females seek the mid- to upper canopy of
the larval food plant for mate location.

Females were usually observed on or near the larval
food plant. Their flight was considerably slower than that
of males but, like males, they sometimes settled on the
outer foliage of the food plant, with wings opened up to
90° towards the sun. They were observed ovipositing from
late morning to early afternoon (1000-1415 hrs EST) during
warm, sunny weather. Multiple females were occasionally
observed to oviposit simultaneously on the same tree.

Life cycle. Adults have been collected from December
to February. At several locations, early instar larvae were
found in early December which, given the developmental
time of the immature stages (see below), indicates that
females were ovipositing in November. Thus, the putative

flight period of Candalides absimilis edwardsi ssp. nov.
is from late November to late February, which coincides
with the period of seasonal flushes of new growth of the
larval food plant. In captivity, eggs hatched about a week
after being laid, and the larvae completed development
in about 3.5 weeks. Pupal developmental time was more
variable: from a sample of 52 pupae, most (81%) reared
from larvae collected in December and January developed
directly, with the pupal duration lasting 11-14 days, but
the others (19%) remained dormant for two months or for
6-10 months, indicating presence of a facultative diapause.
The life cycle, from egg to adult for directly developing
individuals, was completed in approximately six weeks.
These observations suggest that most of the population of
the subspecies is bivoltine, with at least two overlapping
generations during the flight season.

Conservation status. Candalides absimilis edwardsi
ssp. nov. has a narrow geographical range, with an estimated
extent of occurrence of 3,200 km 2 . It is known from eight
sites (> 1 km apart) representing five locations (>10 km
apart) in three disjunct areas. Although the taxon is known
from a limited area, there is no evidence of decline, recent
fragmentation of populations, or threatening processes
impacting on the known sites or habitat, and all known
populations occur in conservation reserves. Its conservation
status should therefore be regarded as Least Concern (LC)
according to IUCN criteria. At Buchan Caves, the low
number of trees of Brachychiton populneus suggests the
butterfly population may be small, and further planting
of B. populneus is recommended to augment the existing
population.

Dunn et al. ( 1994, pp. 236-237) provisionally regarded
Candalides absimilis edwardsi ssp. nov. as secure on the
basis that the known site was protected within a single
national park, but cautioned that the population “...may
become threatened because of their perching behaviour on
bushes along the escaqrment where adults could be taken
in large numbers by hand or with a short handled net.” and
that “Regular wildfires through the main breeding colony
could be a threat and females appear to be associated with
remnant warm temperate rainforest patches which have
escaped burning.” Both of these statements in relation to
threatening processes must be seriously challenged. First,
the breeding area at Mitchell River, based on the extent
and abundance of the larval food plant, is concentrated
along the steep rocky fire-prone escarpment between the
rainforest gully and woodland plateau. Moreover, not
all adults perch along the escarpment, and there is little
documented evidence that butterfly collecting per se is
a threatening process, except perhaps in those situations
where populations have already been substantially reduced
and fragmented by habitat change (New 1991). Second,
the bivoltine life cycle and behaviour of larvae, which
undoubtedly leave the food plant to pupate in concealed
situations near or below the ground, suggests the species
would be well protected from fire.

M. F. Braby

Candalides margaritagilberti Waterhouse,

1903 stat. rev.

(Figs 19-22, 24, 25,29,30)

Candalides gilberti Waterhouse, 1903a, p. 181;
Waterhouse 1903b, p. 23; Waterhouse and Lyell 1914, pp.
78-79, pi. 15; Waterhouse 1932, p. 129, pi. 19; Common
1964, p. 122; Peters, 1971, p. 30; Common and Waterhouse
1972, p. 413, pi. 40; Common and Waterhouse 1981, pp.
527-528, pi. 47; Common and Waterhouse 1982, pp. 301-
302, pi. 26; Puccetti 1991, p. 144, 146; Dunn and Dunn
1991, pp. 389-390; Samson and Wilson 1995, pp. 71-73;
Edwards 1996, p. 252; Daniels and Edwards 1998, p. 90;
Braby 2000, pp. 757-758, pi. 54; Edwards et al. 2001, p.
139; Braby 2004, p. 258; Meyer et al. 2006, p. 13; Franklin
2007, pp. 12-13.

Holochila gilberti (Waterhouse). - Tite 1963, p. 205;
Tindale 1965, p. 173; McCubbin 1971, p.71; D’Abrera
1971, pp. 366-367.

Comments on synonymy. Holochila margarita was
described by Semper (1879) based on an unspecified
number of specimens, but he did not designate a holotype.
Gabriel (1932) gave the type locality as Bowen, Queensland,
and he appears to be the first author to have referred to a
type. Edwards et al. (2001) interpreted Gabriel’s (1932)
reference to a ‘holotype’ as a lectotype designation. The
lectotype male is illustrated in Figures 15-16 and 23; a
typical example of the female of Candalides margarita
margarita (Figs 17, 18) is also illustrated for comparison.
C. margarita gilberti was originally described as a distinct
species by Waterhouse (1903a) based on a pair of specimens
(Figs 19-22) collected from Darwin (given as Port Darwin),
Northern Territory, by Gilbert Turner. Although both
syntypes in AMS possess Waterhouse’s labels indicating
they are type specimens (Figs 24,25), Waterhouse (1903a)
did not designate a holotype. Both specimens were
subsequently illustrated by Waterhouse and Lyell (1914, p.
79) who noted that “The figures arc drawn from the types,
both of which are in poor condition.” Tindale (1965, p.

173) remarked that “The type is from Port Darwin...”, but
did not refer to a holotype. Peters (1971) referred to the
syntype male as the ‘holotype’ and the syntype female as
the ‘allotype’. Edwards et al. (2001) interpreted Tindale’s
(1965) reference to a type as a lectotype designation, but
this interpretation must be rejected since it is not clear
which of the two syntypes Tindale was referring to. In this
work, Peters (1971) incorrect reference to a holotype is
regarded as a lectotype designation according to Article
74 of the 1CZN (International Commission of Zoological
Nomenclature 1999). In his extensive revision of the genus,
Tite (1963) regarded C. maria Bethune-Baker, 1908 from
Waigeo, Misool, Aru and mainland New Guinea to be the
Papuan subspecies of C. margarita based on similarities
in the male genitalia, and this arrangement was followed
by Parsons (1998).

Waterhouse (1903a) drew attention to the distinct
underside markings, especially the postmedian series, and

the reduced white areas on the upperside of the female by
which Candalides margarita gilberti was distinguished.
He considered C. margarita gilberti to be most closely
allied to C. absimilis, but did not compare the taxon with C.
margarita maigarita. Tite (1963) considered C. margarita
gilberti to be specifically distinct based on differences in
the male upperside ground colour, shape of the fore wing
apex, and extent of black spots on the underside of the hind
wing, with C. maigarita gilberti being pale lavender-blue,
having the fore wing apex noticeably produced, and with
sharp black spots on the underside. He also drew attention
to the distinct upperside colouration of the female, which
is pale lavender-blue with the fore wing costal margin
broadly black, but without darkened veins. Tite (1963,
p. 205) regarded C. margarita gilberti to be closely
related to C. margarita, C. tringa (Grose-Smith, 1894)
and C. biaka (Tite, 1963) on account of similarities of the
male genitalia, but remarked that “Surprisingly.. .the male
genitalia arc identical with those of maigarita.'” Parsons
(1986), however, considered C. maigarita sensu stricto
to be closely allied to C. afretta Parsons, 1986 from the
lowland savanna belt of the Western Province of Papua New
Guinea, but he did not refer to C. maigarita gilberti. Braby
(2000, pp. 757-756) stated that “C. gilberti is very closely
related to, and possibly conspccific with, C. margarita ” and
recommended that “Further study is needed to determine if
C. gilberti is merely a subspecies of C. maigarita.”

Type material. LECTOTYPE - cf labelled “Port
Darwin, 24 Nov. 1902, G. Turner | L2008”, “C. gilberti cf.
Type specimen | KL21869”, “G.A. Waterhouse Collection”,
“FIG. 266 Upperside IN ‘THE BUTTERFLIES OF
AUSTRALIA', by WATERHOUSE & LYELL, was taken
from this specimen | KL21869”, “K191348” (AMS).
PARALECTOTYPE - 9 labelled “Port Darwin, 25 Nov.
1902, G. Turner | L2009”, “C. gilberti 9 , Type specimen
| KL2I870”, “G.A. Waterhouse Collection”, “FIG. 267
Upperside IN ‘THE BUTTERFLIES OF AUSTRALIA’, by
WATERHOUSE & LYELL, was taken from this specimen
| KL21870”, “K191349” (AMS).

Other material examined. 58cfcf, 3999. WESTERN
AUSTRALIA: 1 cf labelled “16.31S 125.16E, Synnot Ck
W.A., 17-20 Jun. 1988, T.A. Weir” (ANIC); 19 labelled
“14.25S 126.38E 12 km S of Kalumburu Mission W.A.,
7-11 June 1988, T.A. Weir” (ANIC). NORTHERN
TERRITORY: 19 labelled “48 mi. SW. of Daly River, NT.

14.11S, 130.08E, 3 Sept. 1968, M. Mendum” (ANIC); 4eftf
labelled “60 km S of Daly River N.T., 12 June 1981, D.P.
Sands” (ANIC); 1 9 labelled “DALY R N.T., 30 JUNE
1969, JC LE SOUEF” (ANIC); I cf labelled “16°07’39”S,

130° 19’00”E, Gregory NP, NT, 31 JUL. 2006, M.F. Braby”,
“NT Museum 1004226” (NTM); 1 cf labelled “N. Territory,
BunellsTrig., I3°30’S, 131°02’E, 244m, 23 July 1983, D.P.
Sands” (ANIC); 1 cf labelled “Mt Burrill [sic] NT, 18 Jan
1992, R.N. STOODLEY”, “L.R. Ring Collection” (ANIC);

1 cf labelled “13°29’46”S, 131 °02’08”E, Burrells Trig, NT
250m. 6 MAY 2006, M.F. Braby & DA Young”, “MFB

Taxonomic review of Candalides

Collection 00175” (MFBC); 1 cf labelled similarly but with
registration number “MFB Collection 00083” (MFBC);
3cfcf, 299 labelled “P. Darwin, F.P. Dodd | 4918-4922”,
“KL21874”, “G.A. Waterhouse Collection” (AMS); 19
labelled “P. Darwin, 13 Sep. 08, F.P. Dodd”, "Passed
through C.W. Wyatt Theft-Coil. 1946-1947”, “LEP-9746”
(NMV); 19 labelled “P. Darwin, Nov. 08, F.P. Dodd”,
“Passed through C.W. Wyatt Theft-Coil. 1946-1947”,
“LEP-9745” (NMV); 19 labelled “P. Darwin, Bd. Nov 08,
F.P. Dodd | 4349”, “KL21871”, “G.A. Waterhouse
Collection”, pupal exuvia mounted separately adjacent to
specimen and labelled similarly (AMS); 19 labelled “P.
Darwin, Bred Jan 09, F.P. Dodd”, “Passed through C.W.
Wyatt Theft-Coil. 1946-1947”, pupal exuvia mounted
separately adjacent to specimen and labelled similarly
(AMS); 1 cf labelled“P. Darwin, Jan 09, F.P. Dodd”, “From
F.P. Dodd, N.Q. Land, 12.12.10”,“LEP-9739”(NMV); 1 cf
labelled “P. Darwin, Feb. 09, F.P. Dodd”, “LEP-9740”
(NMV); 19 labelled similarly but with registration number
“LEP-9743” (NMV); 19 labelled similarly but with
registration number “LEP9-747” (NMV); 2cfcf, 19 labelled
similarly but with additional labels “KL21872”, “G.A.
Waterhouse Collection” (AMS); lcf labelled similarly
(ANIC); 19 labelled similarly but with additional label
“Figured in “AUSTRALIAN BUTTERFLIES” Jacaranda
Press, 1964, I.F.B. Common” (ANIC); 1 cf labelled “P.
Darwin, Mch 09, F.P. Dodd”, “Passed through C.W. Wyatt
Theft-Coil. 1946-1947” (AMS); 1 cf labelled “P. Darwin,
Apr. 09, F.P. Dodd”, "Passed through C.W. Wyatt Theft-
Coil. 1946-1947”, “LEP-9741” (NMV); 19 labelled “P.
Darwin, Apl 09, F.P. Dodd | 4464”, “KL21873”, “G.A.
Waterhouse Collection” (AMS); 1 cf labelled “P. Darwin,
May 09, F.P. Dodd”, “Passed through C.W. Wyatt Theft-
Coil. 1946-1947”, “LEP-9742” (NMV); 1 cf labelled “P.
Darwin, May 09, F.P. Dodd”, “LEP-9737” (NMV); lcf
labelled similarly but with registration number "LEP-9738”
(NMV); 19 labelled similarly but with registration number
“LEP-9744” (NMV); 19 labelled similarly but with
registration number “LEP-36622”, “Collection A.N. Bums”
(NMV); 19 labelled “Darwin, Nov., Purcell", “Label data
very doubtful, 22.10.68 J.V.P.”, "Passed through C.W. Wyatt
Theft-Coil. 1946-1947” (AMS); 19 labelled “Darwin NT,
Jan 1949”, “F.E. Parsons Collection, Donated A.N.l.C.
1967” (ANIC); 1 cf labelled “AUSTRALIA, NT DARWIN,
Stuart Pk, T. FENNER ex D. Wilson”, “larva on Decaisnina
signata. cm. 31 May 91” (ANIC); 1 cf labelled “Stuart Park
N.T., 18 June 1991, D.N. WILSON”, “NORTHERN
TERRITORY MUSEUM, 1003598 LYCAEN1DAE,
Candalidesgilberti, Waterhouse, 1903, Det; D.N. Wilson"
(NTM); 19 labelled “Stuart Park N.T., Ex pupa 24 June
1991, D.N. WILSON”, “NORTHERN TERRITORY
MUSEUM, 1003599 LYCAENIDAE, Candalides gilberti,
Waterhouse, 1903, Det: D.N. Wilson” (NTM); 19 labelled
“Stuart Park N.T., Ex pupa 24 June 1991, D.N. Wilson”,
“G. Daniels Collection” (AMS); 299 labelled “Stuart Park
N.T., Ex pupa 25 June 1991, D.N. Wilson” (ANIC); 19

labelled similarly but with additional label “Figured in
Butterflies of Australia (1999) [sic], CSIRO Publishing,

M. F. Braby” (ANIC); 19 labelled “Egg collected 13 May
1991, Egg hatched 15 May 1991, Larvae pupated 31 May
1991, Adult emerged 10 July 1991, Nudle Street Stuart Park

N. T., D.N. WILSON”, “Figured in Butterflies of Australia
(1999) [sic], CSIRO Publishing, M.F. Braby” (ANIC); lcf
labelled “Stuart Park NT, 07 Apr 1992, D.N. Wilson”, “G.
Daniels Collection” (AMS); 19 labelled "NIGHTCL1FF,
NT, 23 AUG 92, C. M EYER XP”, “G. Daniels Collection”
(AMS); lcf labelled “NIGHTCLIFF, NT, 3 SEP 92, C.
MEYER XP”, “G. Daniels Collection”, pupal exuvia pinned
beneath specimen (AMS); lcf labelled “Darwin, N.T.,
Coconut Grove, 2 OCT. 1994 B., C. Meyer". “MFB
Collection 00079”, “genitalia No. 014” (MFBC); lcf
labelled “Fannie Bay, Darwin, N.T., R.P. WeirXP, 3.8.98”,
“NORTHERN TERRITORY MUSEUM, 1003481
LYCAENIDAE, Candalides gilberti, Waterhouse, 1903,
Det: D.F. Trembath 16 Mar 2006” (NTM); lcf labelled
“Darwin, NT. emg. 28 FEB. 2005, M.F. Braby & R. Weir,
The Gardens”, “reared from larva on flowers of Decaisnina
signata, coll. 2 FEB. 2005”, “MFB Collection 00174”,
“genitalia No. 023” (MFBC); 1 9 labelled “Darwin, NT,
emg. 28 FEB. 2005, M.F. Braby & R. Weir, The Gardens”,
“reared from larva on flowers of Decaisnina signata, coll.
2 FEB. 2005”, “MFB Collection 00176” (MFBC); 19
labelled similarly but with date “emg. 8 MAR. 2005” and
registration number “MFB Collection 00080”, pupal exuvia
pinned beneath specimen (MFBC); 19 labelled “Stuart
Park. Darwin, NT, emg. 26 MAY 2006, D.A. Young, reared
from larva on Decaisnina signata", “MFB Collection
00178” (MFBC); 1 cf labelled similarly but with date “emg.
20 MAY 2006” and registration number “MFB Collection
00081” (MFBC); lcf labelled “12°22’S, 130°53’E, Leanyer,
Darwin, NT, emg. 26 JUL. 2006, M.F. Braby”, “reared from
larva on Decaisnina signata, coll. JUL. 2006”, “MFB
Collection 00171” (MFBC); 19 labelled similarly but with
date “emg. 31 JUL. 2006” and registration number “MFB
Collection 00177” (MFBC); 1 cf labelled similarly but with
registration number “MFB Collection 00082” (MFBC); 19
labelled similarly but with date “emg. 2 NOV. 2007”,
“reared from larva on foliage of Decaisnina signata, coll.
OCT. 2007” and registration number “MFB Collection
00179” (MFBC); 19 labelled similarly but with date “emg.
4 NOV. 2007” and registration number “MFB Collection
00180” (MFBC); 1 cf labelled similarly but with registration
number “MFB Collection 00172” (MFBC); lcf labelled
similarly but with date “emg. 24 APR. 2008”, “Reared from
larva on foliage of Decaisnina signata, coll. APR. 2008”
and registration number “MFB Collection 00187” (MFBC);
lcf labelled“12.37308°S, 130.88657°E, Wanguri, Darwin,
NT, emg. 10 FEB. 2007, M.F. Braby”, “reared from egg on
Decaisnina signata, coll. 12 JAN. 2007”, “MFB Collection
00173”, pupal exuvia pinned beneath specimen (MFBC);
19 labelled “21 August, 1970., J. V. Peters, Howard Springs,
N.T., 16 mis, South Darwin.” (AMS); lcf labelled “Tidy

M. F. Braby

Hill 233’ N.T., 12°45’S. 130*54’E., 25 August 1970., J.V.
Peters” (AMS); lcf labelled "13.21S 131.08E, Hill above
Robin Falls N.T., 8 June 1993, E.D. Edwards” (AN1C); 19
labelled “Marrakai Rd, N.T., R.P. Weir XP, 26.10.97”,
“NORTHERN TERRITORY MUSEUM, 1003480
LYCAENIDAE, Candalides gilberti, Waterhouse, 1903,
Det: D.F. Trembath 16 Mar 2006” (NTM); 1 cf labelled “20
KM STH ADELAIDE R. NT., 23 FEB. 1992, RN
STOODLEY”, “L.R. Ring Collection” (ANIC); lcf labelled
“30 KM STH PINE CK NT, 14 APRIL 1992, D.N.
WILSON”, “Figured in Butterflies of Australia (1999) [sic],
CSIRO Publishing, M.F. Braby” (ANIC); 19 labelled
“12.35S 131.17E, Fogg Dam N.T., 15 June 1981, D.P.
Sands” (ANIC); lcf labelled similarly but with date “17
June 1981” (ANIC); 2cfcf labelled “Mt. Bundey - Mary R.
District. N.T., 14 May, 1972. J. Kerr” (ANIC); 1 9 labelled
“11.09S 132.09E, Black Point Cobourg Pen., NT, 26 Jan.
1977, E.D. Edwards” (ANIC); 19 labelled “3 mi. S. of
Oenpelli, NT., 11 Dec. 1970, W. Omer Cooper” (AMS);
lcf labelled “South Alligator RiverN.T., 15 May 1981, D.P.
Sands” (ANIC); 1 cf labelled “Little Menagerie Rock, N.T.,
16 May 1981, D.P. Sands” (ANIC); lcf labelled “12.48S
132.42E, Nourlangie Creek, 8 km N of Mt. Cahill, N.T., 19
Nov. 1972, E.D. Edwards & M.S. Upton” (ANIC); lcf
labelled “12.50S 132.5 IE, 15km NE of Mt. Cahill, N.T.,
23 May 1973, E.D. Edwards & M.S. Upton" (ANIC); lcf
labelled “12.86488°S, 132.70468°E, Mt Cahill (Mirrai
lookout) Kakadu NP, NT, 3 MAY 2008, MF Braby & LJ
Aitchison”, registration number “MFB Collection 00186”
(MFBC); lcf labelled 12°39’19”S, 135°51’31”E, Dbamiyaka
outstation, NE Arnhem Land, NT, 14 AUG. 2007, M.F.
Braby & I. Morris”, “NT MUSEUM 1004303” (NTM); I cf
labelled “Hideway Motel, Gove, 17.iv.l976,A.J. DartncU”,
“NORTHERN TERRITORY MUSEUM, 1003479
LYCAENIDAE, Candalides gilberti , Waterhouse, 1903,
Det: DF Trembath 16 Mar 2006” (NTM); 19 labelled
“12°10’41”S, 136°47’01 ”E, Nhulunbuy, NT, emg. 12 FEB.
2008, MF Braby & LJ Aitchison”, “Reared from larva on
Decaisninasignata, coll. 27 JAN. 2008”; “MFB Collection
00181”, pupal exuvia pinned beneath specimen (MFBC).
QUEENSLAND: 3cfcf, 299 labelled “18.35S 138.03E,
Murray’s Spring 8 km W by N of Musselbrook Camp Q, 9
May 1995 GPS, E.D. Edwards” (ANIC); 3cfcf labelled
similarly but with date “11 May 1995” (ANIC); lcf labelled
“18.38S 138.08E, Gorge 2 km S of Musselbrook Camp Q,
20 May 1995 GPS, E.D. Edwards” (ANIC); lcf labelled
“18.42S 138.29E, Lawn Hill Ck, Q, 17 May 1995 GPS,
E.D. Edwards” (ANIC).

Comparison. Adults of Candalides margarita gilberti
are very similar to those of C. maigarita margarita in wing
shape and pattern, although in C. margarita gilberti the
underside markings are more distinct, and the fore wing
has a straighter termen with the apex more pointed. The
upperside ground colour of males of the two subspecies
is similar, but in C. maigarita gilberti the colouration is
somewhat paler and the black margin is narrower than in

C. maigarita maigarita. C. maigarita gilberti females have
narrower black margins with the large white central patches
on the upperside replaced with blue. A characteristic feature
of C. maigarita gilberti noted by Waterhouse (1903a), that
the postmedian spot between veins Rs and M, displaced
proximally from the remainder of the postmedian series
on the underside of the hind wing, is also shared with
C. margarita margarita but not by other members of the
C. absimilis species group from Australia.

Remarks. The structure of the male genitalia of
Candalides maigarita , which is substantially different and
more complex than those of C. absimilis , is remarkably
uniform across the species range (Figs 29-32). Although the
genitalia are slightly smaller from the Top End and western
Gulf Country (C. maigarita gilberti) compared with those
of from the Wet Tropics and north-eastern New South
Wales (C. maigarita maigarita), there are no significant
differences in the form of the valvae or brachia. The female
genitalia (not illustrated) show a similar geographical
pattern to that of the males.

Larvae of both Candalides margarita gilberti and
C. maigarita margarita specialise on loranthaceous food
plants (Smales and Ledward 1943; Samson and Wilson
1995), and comparison of the immature stages of the two
taxa indicate that the morphology of the egg (Figs 43,44),
final instar larva (Figs 50, 51, 54, 55) and pupa (Figs 52,
53,56,57) are identical. Comparison of the first instar larva
of the two taxa also revealed no differences. The first instar
larva is characterised by a pronounced dorsal ridge along the
length of the body, which bears a series of long paired dorsal
primary setae on the metathorax and abdominal segments
I -7; on each segment lies a pair of smaller setae posterior
to the longer paired setae. The mesothorax also possesses
a pair of long setae, but they are located dorsolaterally so
that they are not as close together as the setae on the other
segments. Final instar larvae of the two subspecies vary
in pattern and colouration depending on the colour of the
food consumed (red flowers or green leaves), although in
C. maigarita gilberti the darker patches on the body appear
to be more pronounced.

In summary, evidence from the genitalia, adult and
immature stage morphology, together with similarities in
the life history and adult behaviour (see below), indicate
that gilberti is conspecific with Candalides margarita ;
gilberti should be regarded as a geographical subspecies
of that taxon on account of its disjunct distribution and
morphological divergence in the adult stage. Thus, three
subspecies are now recognised within C. margarita :
C. maigarita maria from mainland New Guinea and the
nearby islands of eastern Indonesia; C. maigarita gilberti
from north-western and central northern Australia; and
C. maigarita maigarita from eastern Australia (Thursday
Island, Queensland, to Tuncurry, New South Wales).
Presumably, C. margarita gilberti is sister to C. margarita
margarita + C. margarita maria based on the greater
phenotypic divergence of C. maigarita gilberti and the close

Taxonomic review of Candalides

similarity among the females of C. margarita margarita
and C. margarita maria.

Distribution. Candalides margarita gilberti occurs in
the monsoon tropics of north-western and central northern
Australia, from the Kimberley, Western Australia, across
the Top End, Northern Territory, to the Gulf Country of
north-western Queensland (Braby 2000; Franklin 2007). It
is allopatric from C. margarita margarita, the known ranges
of the two subspecies in Queensland being separated by a
minimum distance of about 700 km. C. margarita gilberti
reaches its northernmost limit at Black Point on Cobourg
Peninsula (Garig Gunak Barlu National Park), Northern
Territory, about 550 km south-south-west of Aru Island in
the Arafura Sea which supports the nearest population of
C. margarita maria.

Habitat. Candalides margarita gilberti, in contrast
to C. margarita margarita, occurs in savanna woodland
and patches of monsoon forest and vine-thicket where the
larval food plant grows. In Darwin, it is widespread and
the early stages may be relatively common in suburban
areas, parklands and areas with native vegetation where
the subspecies breeds throughout the year.

Biology. The larval food plant, description of the
early stages, general biology and habitat of Candalides
margarita gilberti have been well documented (Samson
and Wilson 1995; Braby 2000). The recorded larval food
plant is the aerial-stem hemiparasite Decaisnina signata
(F.Muell. ex Benth.) Tiegh. (Loranthaceae) (Samson and
Wilson 1995), but eggs have also been found on Decaisnina
petiolata (Barlow) Barlow at Kakadu National Park (M.F.
Braby unpublished data). Although the life history was
documented only relatively recently, Waterhouse and
Lyell (1914, p. 79) noted earlier that “The species has
since been bred by Mr. F.P. Dodd...” Examination of the
Dodd material collected from Darwin during 1908-09 in
AMS,ANICandNMV(13cfcf, 139 9) revealed two reared
female specimens each with their pupal exuviae mounted
separately and labelled “P. Darwin, Bd. Nov 08, F.P. Dodd”
(AMS) and “P. Darwin, Bred Jan 09, F.P. Dodd” (AMS);
however, no information on the identity of the larval food
plant was provided on the labels of these specimens.

Little has been documented on adult behaviour
of Candalides margarita gilberti, but the following
observations suggest it is similar, if not identical, to that of
C. margarita margarita (M.F. Braby unpublished data). In
the Northern Territory, C. margarita gilberti males regularly
congregate on hilltops where they perch on foliage of trees
to defend territories during the morning (from 7.45 am
onwards) and the afternoon. When settled, they usually
perch, with wings closed, 3-5 m above ground level. Males
also establish territories along edges of gallery forest where
they typically fly and perch, with wings closed and oriented
head downwards towards the sunlight, on outer foliage of
rainforest trees about 5-8 m above ground level during
early to late afternoon. Females have been observed flying
in similar places, but when settled frequently perch with

wings opened at about 90° towards the late afternoon sun.
They have been observed ovipositing on the larval food
plant during the early afternoon.

Conservation status. The conservation status of
Candalides margarita gilberti should be regarded as
Least Concern (LC) on account of its broad distribution
and lack of threatening processes. No populations are
known to have been extirpated and there is no evidence
that the subspecies has declined or is in decline. However,
its relative abundance in savanna woodland across the
monsoon tropics of the Kimberley and Top End may have
declined over the past century with changed fire regime
because mistletoes, the larval food plant of the butterfly,
are highly susceptible to fire (Kelly et al. 1997). In these
fire-prone habitats, there has been a general increase in
the extent, frequency and intensity of fires, in contrast to
the general suppression of fire since European settlement
in agricultural areas of temperate southern Australia
where some mistletoe species appear to have increased in
abundance (Kelly et al. 1997).

DISCUSSION

Geographical variation in Candalides absimilis males
parallels that of C. consimilis males (Tindale 1965; Kerr
1967), both species of which are broadly sympatric along
the eastern coast of Australia. Not only do these species
diverge at the extreme ends of their geographical ranges, but
they also exhibit similar patterns of phenotypic divergence,
particularly on Cape York Peninsula where they both have
narrow range endemic subspecies restricted to the mid¬
peninsula rainforest block (Iron Range-Mcllwraith Range).
Males of both C. absimilis eastwoodi and C. consimilis
toza, which are superficially similar, are characterised
by possessing a paler upperside purple ground colour, a
straightcr termen of the fore wing, a more prominent black
tomal spot on the hind wing, and less distinct markings
on the underside compared with their respective nominate
subspecies. This suggests that the northernmost populations
of the two species of Candalides, C. absimilis eastwoodi
and C. consimilis toza, have diverged under similar selective
pressures in both space and time. The mid-peninsula block
of rainforest on Cape York Peninsula is well known for its
high level of local endemism and insularity (Kikkawa etal.
1981), and the biogeographical patterns observed in the two
species of Candalides suggest these butterflies have been
subjected to the same underlying historical processes that
have led to isolation and differentiation.

Interestingly, differences observed between males of
the two Australian subspecies of Candalides margarita
parallels the pattern of geographical variation observed in
fore wing shape in C. absimilis and C. consimilis males in
which the subspecies from the Kimberley and Top End,
C. margarita gilberti, has the termen straighter and apex
sharply pointed, similar to that observed in C. absimilis

M. F. Braby

eastwoodi and C. consimilis toza. C. margarita gilberti
also has more prominent black tomal spots on the hind
wing underside than in C. margarita margarita, but the
underside basal spots and wavy lines are more distinct,
whereas in C. absimilis eastwoodi and C. consimilis toza
the underside markings are less distinct than their respective
nominate subspecies. In C. margarita, the geographical
variation is partitioned across the ‘Carpentarian Gap’, a
barrier comprising the Gulf of Carpentaria and arid plains
of the Gulf Country between the Top End and Cape York
Peninsula, whereas in C. absimilis and C. consimilis
geographical variation among the northernmost populations
is partitioned across a putative barrier comprising the dry
lowland plains south of Princess Charlotte Bay of Cape
York Peninsula (i.e. between Iron Range-Mcllwraith Range
and the Wet Tropics).

Although the blue colouration differs between females
of Caudal ides absimilis edwardsi and C. margarita gilberti,
the upperside pattern of these two subspecies is broadly
similar in that the white central patches are absent and the
basal and central areas of both wings are broadly blue.
Indeed, these two subspecies are remarkable in that they
comprise the only taxa in the C. absimilis species group
in Australia in which the white patches are replaced with
blue. The only other species in the C. absimilis species
group that possess this trait are C. riuensis (Tite, 1963)
from Tagula (Sudest) Island, C. lamia (Grose-Smith,
1897) from the D’Entrecasteaux Islands, and C. coeruleus
(Rober, 1886) from Aru Island and the lowland areas
(< 200 m) of mainland New Guinea (Parsons 1998).
These taxa, together with C. cyprotus , are exceptional as
females of most members of the C. absimilis species group
have the upperside ground colour black with contrasting
patches of white in the central areas of the wings, and
their morphological pattern poses interesting questions
as to how this character may have evolved, its functional
significance and whether the ‘blue form’ represents a
plesiomorphic (ancestral) or apomorphic (derived) state.
Answers to such questions may come from comparative
analysis of female phenotype in relation to the species’
ecology and phylogenetic history of the genus. For instance,
among the Australian taxa, both C. absimilis edwardsi and
C. margarita gilberti are associated mainly with habitats
comprising eucalypt woodland, open woodland and savanna
woodland (i.e. vegetation types in which the projective
foliage cover < 30%), whereas C. absimilis absimilis, C.
absimilis eastwoodi, C. maigarita margarita, C. helenita
(Semper, [1879]), C. consimilis consimilis and C. consimilis
toza are associated primarily with closed forest habitats,
particularly rainforest (i.e. vegetation types in which the
projective foliage cover > 70%) (Braby 2000). Although C.
margarita gilberti also occurs in dry rainforest (monsoon
forest), this vegetation type in the Kimberley and Top End,
compared with the evergreen tropical forests of eastern
Australia, occurs as a vast network of smaller patches
(mostly < 5 ha in extent) that is naturally fragmented

throughout the extensive savanna landscape so that these
habitats have greater edge effects (i.e. larger perimeter/area
ratios) where the canopy is more open and, depending on
water availability, the dominant canopy trees are frequently
deciduous or semi-deciduous during the dry season.
Thus, the ‘blue forms’ in female Candalides are primarily
associated with open habitats, whereas the striking and
highly contrasting ‘black and white forms’ are associated
with closed habitats. In this context, it is noteworthy that
C. absimilis absimilis females from the extreme southern
end of its range in south-eastern coastal New South Wales
(Central Tilba-Bega), where it breeds primarily in eucalypt
open-forest and tall open-forest (projective foliage cover
30%-70%), have substantially reduced white patches on the
wings (Fig. 34F) compared with populations further north
that arc mainly associated with closed forest. A similar trend
also occurs in C. consimilis goodingi females from south¬
eastern Australia, which occurs primarily in eucalypt tall
open-forest, in which the white patch on the fore wing is
reduced in extent compared with the nominate subspecies
C. consimilis consimilis, which occurs mainly in subtropical
and warm temperate rainforest.

The association between sex-limited phenotype and
habitat (i.e. broad vegetation type, measured in terms of
projective foliage cover) in Candalides also extends to other
species in the family from Australia (Table 1). Of the 49
species of Lycaenidae that occur predominantly in closed
forest vegetation types, 30(61 %) have the upperside ground
colour black with prominent white patches in the female sex
and the sexes are frequently strongly sexually dimorphic,
whereas of the 90 species that live primarily in open forest/
woodland vegetation types the proportion of ‘black and
white forms’ among females drops to 3%. The three species
in the latter category include two [ Hypolycaena phorbas
(Fabricius, 1793), Erysichton palmyra (C. Felder, 1860)]
which also occur in rainforest and one [Leptotes plinius
(Fabricius, 1793)] in which the white patches are very
obscure. Differences in the frequencies of adult female
forms among the two broad vegetation types tabulated in
Table 1 are significant (%= 55.8, d.f 1, P<0.001), supporting

Tabic 1 . Associations between female phenotype (upperside wing
pattern) and habitat (broad vegetation type) among Australian
Lycaenidae (data compiled from Braby 2000). Differences in
frequency of female forms among the two vegetation types are highly
significant (P < 0.001).

Habitat

Phenotype

Total

Ground colour
black, with
white patches
present*

Ground colour
variable, but
white patches
absentf

Closed forest

Open forest/
woodland

Total

106

139

* white patches may have some traces of blue pigmentation,
t ground colour frequently brown or blue.

Taxonomic review of Candalides

the hypothesis that habitat characteristics affects female
wing phenotype in Candalides, at least in terms of the
visible spectrum. Further analysis is needed to establish
if other factors, such as phylogenetic history and mate
location or ovipositing behaviour, are correlated with
the observed pattern, and if the relationship also holds in
the non-visible spectrum such as ultraviolet light, which
butterflies frequently use for mate recognition. Female
wing pattern does not appear to be explained by shared
ancestry because the trend for habitat associated sex-limited
phenotype among the Australian Lycaenidae also occurs in
Jamides and Nacaduba, two genera which occur in both
open and closed forest.

The evolutionary forces that may have selected for
female wing pattern among Candalides, and the Australian
Lycaenidae in general, have not been established, but since
the two habitats (broad vegetation types) analysed differ
greatly in projective foliage cover and therefore their
light environment, differences in ambient light properties
may be a critical factor. Closed forest environments
are optically complex, varying greatly in intensity
(brightness) and spectral composition (colour) under sunny
conditions (Endler 1992, 1993). Therefore, selection is
expected to favour phenotypes that are more striking to
maximise brightness contrast and conspicuousness among
conspecifics or potential mates. A recent study (Douglas
et al. 2007) has demonstrated that nymphalid butterflies
inhabiting understorey microhabitats of tropical forest, in
which light intensity is very low, are more likely to exploit
polarised light as a signal for communication than related
species occupying open habitats. In contrast, in open
forest/woodland habitats or seasonally deciduous monsoon
forests, Endler (1992) predicted that selection should favour
exploitation of the shorter wavelengths (ultraviolet, blue,
green) for visual signals in these ‘woodland shade’ light
environments. These findings suggest that a more general
hypothesis may be proposed, namely that ambient light
characteristics among divergent habitats or vegetation
types (i.e. different light environments) is a potent selective
force in shaping sex-limited phenotype among Australian
lycaenid butterflies.

ACKNOWLEDGMENTS

I am grateful to D.R. Britton and S.F. McEvey (AMS),
E.D. Edwards and V. Rangsi (ANIC), B. Done (MTQ), R
Lillywhite and C. McPhee (NMV) and J. Forrest (SAM)
for access to and/or loan of specimen material held in their
care. R.G. Eastwood, S.F. McEvey, R.P. Field and C.E.
Meyer also provided material and field notes from their
collections for detailed examination. B. Huertas (BMNH)
kindly provided digital images of type material lodged in
the Natural History Museum, London, for inclusion in this
work, D. Schmidt provided live material of the early stages
of C. margarita margarita from Brisbane for comparative
morphological work, D.J. Kemp greatly assisted with

literature and ideas on animal communication in relation
to light environments, and R.P. Weir introduced me to the
life history of C. margarita gilberti in Darwin. I thank L.J.
Aitchison for field assistance in south-eastern Australia,
and R.P. Field, D.A. Lane and G. Guy for biological
observations on C. absimilis. D. Schmidt and C.J. Muller
provided constructive editorial suggestions and thought-
provoking comments on the manuscript. Specimens of
C. absimilis edwardsi were collected under Research Permit
Number 10002433 issued by the Victorian Department of
Sustainability and Environment.

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Accepted 11 July 2008.

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 55-61

Two new species of the genus Pseudaptinus Castelnau from northern Australia
(Insecta: Coleoptera: Carabidae: Zuphiinae)

MARTIN BAEHR

Zoologische Staatssammlung Miinchen, Miinchhausenstr. 21, D-81247 Miinchen, GERMANY

martin.baehr@zsm.mwn.de

ABSTRACT

Two new species of the zuphiine genus Pseudaptinus Castelnau are described from the Northern Territory of Australia:

P. parallelus sp. nov. and P. gregoryensis sp. nov. According to the size of eye and the structure of surface of the elytra,
the first species is probably most closely related to P punctatostriatus Bachr and P. cyclophthalmus Baehr, but il is
smaller than either of them; the second species is probably nearest to P. iridescens Baehr, but it is distinguished by its
smaller eyes and shorter antennae. Additional records are provided for P. hirsutulus Baehr and P. iridescens Baehr. A
new key to all the Australian species of this genus is included.

Keywords: Coleoptera, Carabidae, Zuphiinae, new species, Pseudaptinus parallelus, P. gregoryensis, taxonomy, Australia,
Northern Territory.

INTRODUCTION

During a recent collecting trip by the author through
the northern parts of the Northern Territory and Western
Australia, two new species of the zuphiine genus
Pseudaptinus Castelnau were collected. These are described
here as P. parallelus sp. nov. and P. gregoryensis, sp.
nov., and this is the fifth supplement to my revision of the
Australian Zuphiinae (Baehr 1984, 1985a, 1985b, 1986a,
1985b, 1988, 1991, 1992, 1995, 2001, 2008). Additional
specimens of one of the new species, P. parallelus,
were also received from the Australian National Insect
Collection, Canberra, and the South Australian Museum,
Adelaide. Additional records of two other species of this
genus occurring in northern Australia, P. hirsutulus Baehr
and P. iridescens Baehr, are also provided. The key to the
11 Australian species of this genus is revised to include all
species described since the revision of the genus by Baehr
(1985).

With about 40,000 described species and subspecies
worldwide, carabid beetles are one of the largest families
of beetles. They belong to the suborder Adephaga which
also includes a few families of aquatic beetles and is mainly
characterised by a rather plesiomorphic structure of their
leg insertions and by their primitive campodeiform larvae
which still possess well articulated legs. From Australia
presently about 2,600 species and subspecies are recorded,
but the number of actual existing species certainly is much
greater.

Zuphiinae (or Zuphiini, according to the opinion of
authors) is a highly evolved subfamily (or tribe) of carabid
beetles, characterised by the completely pilose antennae and
in particular by the elongate 1st antennomere, the scapus.

Opinions about the status (subfamily or tribe) of the main
subgroups of the family Carabidae differ markedly between
authors, but the delimitation of the subgroups is not in doubt,
hence the question whether zuphiines should be regarded a
subfamily or rather a tribe, is of minor importance.

Zuphiinae presently contain eight genera in Australia,
of which the genus Acrogenys Macleay and two genera
of cavernicolous species ( Speothalpius Moore and
Speozuphium Moore) are endemic to Australia. With respect
to their phylogenetic status, the Australian zuphiine fauna
is particularly diverse and includes as well the probably
most plesiotypic zuphiine genus in the world ( Acrogenys),
as well as the highly evolved genera Speothalpius and
Speozuphium.

At present, 45 zuphiine species and subspecies are
recorded from Australia (Baehr 1984,1985a, 1985b, 1986a,
1986b, 1988, 1991, 1992,1995,2001,2008; Moore 1995),
but specimens of most species still are rare to very rarely
encountered. This is probably due to the presumably rather
secretive way of life of most species which, at least during
daytime, may be hidden under debris or in cracks in the
earth. As a consequence, most recorded specimens were
collected using a light at night.

The genus Pseudaptinus was founded by Castelnau
(1835) on an American species. Later Castelnau (1867)
described a single Australian species which was included
in the subgenus Thalpius Leconte, 1851. Baehr (1985)
revised the genus, transferred Acrogenys australis
Blackburn, 1890 to Pseudaptinus, and described six new
species. Later Baehr (1995) described another additional
species. Hence, at present nine species are recorded from
Australia, all are included in the subgenus Thalpius,
and most of them occur in the tropical northern parts of

M. Baehr

Australia. The genus Pseudaptimis is characterised by the
quite compact build of all species with fairly convex dorsal
surface, their posteriorly rounded, not square head, the only
moderately elongated 1 st antennomere, and the erect and
hirsute pilosity. The subgenus Thalpius is rather weakly
characterised, distinguished from the nominate subgenus
mainly by the more depressed dorsal surface and the wider,
more cordiform pronotum.

The genus Pseudaptimis also occurs in South America
and in the southern parts of North America, where
specimens of both subgenera are found. Hence this is one
of the few carabid genera with an Australian-Neotropical
range, which is very interesting in the light of their
biogeographical history.

METHODS

For the taxonomic treatment standard methods were
used. The male genitalia were removed from specimens
soaked overnight in a jar under a wet atmosphere, then
cleaned for a short while in hot potassium hydroxide.

For examination of the fine, though taxonomically
important, surface microsculpture (i.e., punctation and
microreticulation) a high resolution stereomicroscope with
up to 64x magnification was used, supported by a lamp of
higli intensity giving natural light that could be focused.
For exact definition of the microsculpture such light is
preferable, because fibre-optics lights substantially change
perception of the surface structures.

The habitus photographs were taken with a digital
camera using SPOT Advanced for Windows 3.5 and
subsequently edited with Corel Photo Paint 11.

Measurements were made using a stereomicroscope
with an ocular micrometer. Length has been measured from
the apex of labrum to the apex of the elytra. Length of eye
includes a small ring of dark coloured rhabdomes that in
some instances is present behind the light area. Length of
orbit was taken in a straight line parallel to midline of body,
from the posterior margin of the eye to the ‘neck' incision.
Length of pronotum was measured from the most advanced
part of base to the most advanced part of apex. Length of
elytra was taken from the most advanced part of humerus
to the most advanced apex of elytra. The length/width ratio
of the 6th antennomere was chosen for better comparison
with the measurements in the revision. It should be noted,
however, that the 7th to 9th antennomcres usually are
slightly shorter than the 6th antennomere.

ABBREVIATIONS

ANIC Australian National Insect Collection, Canberra
CBM Working Collection M. Baehr in Zoologische
Staatssammlung, Miinchen

NTM Museum and Art Gallery of the Northern Territory,

Darwin (formerly Northern Territory Museum)
SAMA South Australian Museum, Adelaide

TAXONOMY

Pseudaptimisparallelus sp. nov.

(Figs 1-3, 5, 7)

Material examined. HOLOTYPE-male, Australia07,
NT22, Roper River Crossing, 3 km E. Roper Bar,
14°42.83'S, 134 o 30.51’E, 10 m, 6-7 November 2007, M.
Baehr (NTM 1005298). PARATYPES - 2 females, same
dataasholotype(CBM); 1 female, Australia07, NT9, Mary
River National Park, Maiy River Billabong, 5 km NNW.
Mary River Crossing, I2°53.49’S, 131°38.33’E, 28 m,
1-2 November 2007, coll. M. Baehr (CBM); 1 male, 1
female, 12°40’S 142°40’E, Queensland, Batavia Downs
Homestead. 22 November 1992, light trap, coll. A. Caldcr
and P. Zborowski (ANIC, CBM); 1 male, Old Greenvale,
70 km SW. at light, 14-23 February 96, coll. A. J. Watts
(SAMA).

Diagnosis. Species of subgenus Thalpius Leconte,
characterised by small size, elongate and remarkably
parallel-sided elytra, and short antenna.

Description. Measurements. Length: 4.75-4.95 mm;
width: 1.5-1.6 mm. Ratios: Length of orbit/lcngth of eye:
0.45-0.5: width/length of pronotum: 0.94-0.98; widest /
narrowest diameter of pronotum: 1.31-1.35; length/width
of elytra: 1.77-1.79; length/width of 6th antennomere:
1.75-1.8.

Colour. Reddish-black to blackish, according to age of
the specimen. In dark specimens, suture of elytra indistinctly
paler. Mouthparts, antennae, and legs dirty yellow.

Head (Fig. 3). Slightly narrower than pronotum. Eyes
large and laterally well protruded, about twice as long as
orbits (when measured in a straight line). Orbits markedly

Fig. I. Pseudaptimis parallelus sp. nov. Male genitalia: aedeagus,
lateral and ventral view, left and right parameres. Scale: 0.25 mm.

New species of Pseudaptimis from northern Australia

Fig. 2. Pseudaptimis parallelus sp. nov. Female gonocoxites 1 and
2. Scale: 0.1 mm.

convex, even obtusely ungulate, with sharp angle to neck.
Suture between occiput and summit deep, neck dorsally
and laterally deeply separated. Orbits separated from
summit by sulcus. Posterior supraorbital seta located at
level with posterior margin of eye. Labrum anteriorly
straight. Mandibles short, incurved at apex. Mental tooth
triangular, short and wide. Glossa corneous, apically
square, paraglossae membraneous, short, fused to glossa,
laterally little surpassing the glossa. Galea distinctly pilose.
Both palpi densely pilose, labial palpus small, terminal
palpomcre cylindrical. Maxillary palpus large, though rather
short (in group). Antenna short and stout, just surpassing
base of elytra, median antennomeres distinctly less than
twice as long as wide. 1 st antennomere barely longer than
2nd and 3rd antennomeres together. Surface glossy, not
microreticulate, coarsely punctate, diameter of punctures
slightly larger than distance between punctures, pilosity
fairly dense, hirsute, slightly inclined anteriad.

Prothorax. Slightly longer than wide, depressed,
widest in apical fifth. Apical angles broadly rounded, apex
faintly concave, lateral margins moderately convex on
anterior half, rather deeply sinuate in front of the acute,

Figs 3,4. Left side of head, showing ratio eye/orbit; 3. Pseudaptimis
parallelus sp. nov.; 4. P. gregoryensis sp. nov.

about rectangular.markedly projecting basal angles. Base
distinctly projecting medially, laterally deeply excised,
hence basal angles removed from base. Median line
deep, anterior transverse sulcus indistinct. Central part
of pronotum slightly raised, pronotum near apex with an
elongate, rather deep impression on either side. Base and
apex both unmargined, lateral borders margined, marginal
channel very narrow. Anterior marginal seta situated at
apical fifth, posterior seta at basal angle. Anterior seta
considerably longer than the posterior seta. Surface without
microreticulation, glossy, densely and rather coarsely
punctate, diameter of punctures wider than distance
between them. Pilosity rather elongate, hirsute, irregularly
inclined posteriorly.

Elytra (Figs 5, 7). Narrow and elongate, remarkably
parallel-sided, not perceptibly widened in apical part,
surface depressed. Humeri projected, rounded. Apex
laterally rounded, in middle transversal, barely incurved
towards suture. All striae well impressed, coarsely punctate.
Intervals depressed, without microreticulation, glossy, with
coarse, irregularly uni- to biseriate punctation. Pilosity
hirsute but rather depressed, inclined posteriorly, hairs
about as long as one interval wide. Third interval with three
punctures and erect setae which are well recognisable when
seen laterally.

Ventral surface. Thorax with coarse and moderately
dense punctures and pilosity. Abdominal sterna with finer
and denser punctures and with dense adpressed pilosity.
Metepisternum elongate, greater than twice as long as
wide at anterior margin. Apical margin of sternum VII in
male bisetose, in female quadrisetose, in both sexes with an
additional seta on either side removed from apex.

Legs. Of normal shape and length, densely punctate and
pilose. Three basal tarsomeres of male protarsus slightly
widened and asymmetrically biseriate'y squamose.

Male genitalia (Fig. 1). Aedeagus small, fairly elongate,
apex rather elongate (in group), with remarkably acute
tip. Lower surface very gently concave. Orificium very
elongate, with a median cleft at apex. Internal sac with two
large, rather symmetrical, slightly sclerotised and somewhat
folded plates on either side. Parameres very dissimilar, both
rounded at apex, right paramere remarkably short.

Female genitalia (Fig. 2). Gonocoxite 2 wide, trapezoidal,
with wide, oblique apex, apical part less sclerotised than
basal-central part and densely setose latero-apically. No
ensiform setae present. Apex of gonocoxite 1 asetose, though
apical margin of lateral plate with a few elongate setae.

Variation. Apart from differences in colour which are
probably due to the age of the specimens, little variation was
noted, even between specimens from Northern Territory
and those from northern Queensland.

Collecting circumstances. The specimens from the
Northern Territory were sampled with a black light near
the sandy bed of the Roper River and at the margin of
a billabong with dense marginal vegetation of trees and
undergrowth.

M. Baehr

Figs 5,6. Habitus. Body lengths in brackets; 5. Pseudaptinus parallelus
sp. nov. (4.85 mm); 6. P. gregoiyensis sp. nov. (5.35 mm).

Distribution. North-eastern part of Northern Territory
and tropical northern Queensland, from Cape York
Peninsula southwards to about Townsville.

Etymology. The name refers to the remarkably parallel¬
sided elytra. It is an adjective.

Relationships. Although similar to P. hirsutulus
Baehr in body shape, the new species is certainly closer
to P. punctatostriatus Baehr and P cyclophthalnuis Baehr,
according to the shape of the pilosity and the structure or
the aedeagus. However, it differs from both these species
by its smaller size, shorter antenna, and the conspicuously
parallel-sided elytra.

Pseudaptinus gregoiyensis sp. nov.

(Figs 4, 6, 8)

Material examined. HOLOTYPE-female,Australia07,
NT37, Gregory National Park, Old Victoria River Crossing,

Figs 7, 8. Centre of elytra showing elytra! microsculpture.
7. Pseudaptinusparallelus sp. nov.; 8. P. gregoiyensis sp. nov.

5 km W. Victoria River Roadhouse, Victoria Highway,
15°34.87’S, 131°06.24'E, 35 m, 11-12 November 2007
coll. M. Baehr (NTM 1005299).

Diagnosis. Characterised by moderately large eyes (in
group), rather dense punctures on the elytra! intervals and
fine punctures on the striae. Distinguished from most similar
P. iridescens Baehr by smaller size, slightly smaller eyes,
distinctly shorter antenna, and moreover by far less distinct
iridescent lustre of the elytra.

Description. Measurements. Length: 5.35 mm; width:

1.75 mm. Ratios: Length of orbit/length of eye: 0.75; width/
length of pronotum: 0.91; widest/narrowest diameter of
pronotum: 1.31; length/width of elytra: 1.69; length/width
of 6th antennomere: 2.05.

Colour. Dark blackish brown to almost black, only
pronotum very slightly paler, and suture of elytra narrowly
dark reddish. Mouth parts, antennae, and legs dirty
yellow.

Head (Fig. 4). Slightly narrower than pronotum. Eyes
of moderate size, little protruded laterally, about 1.5 x as
long as orbits (when measured in a straight line). Orbits
evenly convex, forming sharp angle with neck. Suture
between neck and summit of head deep, neck dorsally
and laterally deeply separated. Orbits separated from
summit by sulcus. Posterior supraorbital seta located
slightly behind posterior margin of eye. Labrum anteriorly
straight. Mandibles short, incurved at apex. Mental tooth
triangular, short and wide. Glossa corneous, apically square,
paraglossae membraneous, short, fused to glossa, laterally
little surpassing the glossa. Galea distinctly pilose. Both
palpi densely pilose, labial palpus small, terminal palpomere
cylindrical. Maxillary palpus large, though rather short (in
group). Antenna rather short and stout, sutpassing base of
elytra by about 2.5 antennomeres, median antennomeres
distinctly only twice as long as wide, 1st antennomere
slightly longer than 2nd and 3rd antennomeres together.
Surface glossy, with extremely superficial microreticulation
which is not visible when seen from above, rather densely
punctate, diameter of punctures about as large as distance
between punctures, but laterally punctation less dense,
pilosity fairly dense, hirsute, slightly inclined anteriad.

Prothorax. Considerably longer than wide, depressed,
widest in apical fifth. Apical angles broadly rounded, apex
faintly concave, lateral margins in anterior half moderately
convex, with elongate sinuation in front of the acute,
about rectangular, markedly projecting basal angles. Base
in middle distinctly projecting, laterally deeply excised,
hence basal angles removed from base. Median line
deep, anterior transverse sulcus indistinct. Central part of
pronotum slightly raised, near apex with an elongate, rather
deep impression on either side. Base and apex both not
margined, marginal channel veiy narrow. Anterior marginal
seta situated at apical fifth of length, posterior seta at basal
angle. Anterior seta considerably longer than posterior
seta. Surface with extremely superficial microreticulation,
visible only in oblique light, glossy, densely and moderately

New species of Pseudaptinus from northern Australia

coarsely punctate, diameter of punctures about as wide or
even slightly wider than distance between them. Pilosity
rather elongate, hirsute, irregularly inclined posteriorly.

Elytra (Figs 6, 8). Rather narrow and elongate, fairly
parallel-sided, slightly widened in apical part, surface
in middle depressed. Humeri projected, rounded. Apex
laterally rounded, in middle slightly oblique, barely
incurved towards suture. All striae well impressed,
moderately coarsely punctate. Intervals depressed, with
superficial traces of slightly transverse microreticulation,
rather glossy, with fairly coarse, irregularly triseriate
punctation. Pilosity hirsute but rather depressed, inclined
posteriad, hairs about as long as one interval wide. Third
interval with three punctures and erect setae which are well
recognisable when seen laterally.

Lower surface. Thorax with coarse and moderately
dense punctures and pilosity. Abdominal sterna with finer
and denser punctures and with dense adpressed pilosity.
Metepistemum elongate, about twice as long as wide at
anterior margin. Apical margin of sternum Vll in female
quadrisetosc and with an additional seta on either side
removed from apex.

Legs. Of normal shape and length, densely punctate and
pilose. Squamosity of male protarsus unknown.

Male genitalia. Unknown.

Female genitalia. Similar to those of P. parallelus sp.
nov.

Variation. Unknown.

Collecting circumstances. Collected at a black light
near the bed of the Victoria River. Here the substrate
consists of pebbles and more or less coarse sand.

Distribution. North-western Northern Territory. Known
only from type locality.

Etymology. The name refers to the collecting area,
Gregory National Park. It is an adjective.

Relationships. The species is probably most closely
related to P. iridescens Baehr which is widely distributed
in tropical northern Australia, but it is distinguished from
P. iridescens mainly by the shorter antenna, smaller eyes,
coarser punctures of the elytral striae, and barely iridescent
elytra.

Pseudaptinus hirsutulus Baehr

Pseudaptinus hirsutulus Baehr, 1985a: 51. - Baehr
1986b: 162; Baehr2001: 85.

Distribution. Northern parts of the Northern Territory
from Darwin eastwards at least to the western boundary of
Arnhem Land.

New records. Australia07, NT9, Mary River National
Park, Mary River Billabong 5 km NW. Mary River
Crossing, 12°53.49’S, I31°38.33’E, 28 m, 1-2 November
2007, coll. M. Baehr (CBM, NTM).

Collecting circumstances. All specimens collected
at a black light at the margin of a billabong with dense
vegetation of trees and undergrowth. At the Maty River

Billabong this species was collected together with P.
parallelus sp. nov. (see above).

Pseudaptinus iridescens Baehr

Pseudaptinus iridescens Baehr, 1985a: 43. - Baehr
1986b: 162; Baehr2001: 84.

Distribution. North-western Victoria, New South Wales
and Queensland west of the Great Dividing Range, northern
parts of Northern Territory and of Western Australia.

New records. NT22, Roper River Crossing, 3 km E.
Roper Bar, 14°42.83’S, 134°30.51’E, 10 m, 6-7 November
2007, coll. M. Baehr; NT29, 70 km N. Top Springs on
Buntine Highway, 16°00.94’S, 131°56.33’E, 253 m, 9-10
November 2007, coll. M. Baehr (CBM, NTM).

Collecting circumstances. All the specimens were
collected at a black light, those near Roper Bar close to
the bank of the Roper River; the one specimen from the
‘Buntine Highway’ was taken in open woodland, apparently
far away from any body of water. At the Roper River this
species was collected together with P. parallelus sp. nov.
(see above).

KEY TO GENUS PSEUDAPTINUS

Because the three additional species must now be
included in the key for the genus that was previously
published (Baehr 1985), a completely new key to all the
species of Pseudaptinus is provided. For the benefit of the
users, figures from previous papers arc included as BA85
and BA95.

1 a. Eyes not longer than orbits (BA85 figs 3a-c, g).... 2
lb. Eyes considerably longer than orbits (BA85 figs
3d-f, h).6

2a. Antenna very elongate, median aniennomeres >3x

longer than wide (BA85 figs 4c, f).3

2b. Antenna shorter, median antennomeres <2.5 x longer
than wide (BA85 figs 4a, b).4

3a. Eyes perceptibly shorter than orbits (BA85 fig. 3g);
pronotum elongate, narrow, basal angles very acute
(BA85 fig. 12); pronotum and elytra rather convex;
striae very coarsely punctate, punctures of intervals in
two rows (BA85 fig. 5g); pilosity unequal, rather erect;
aedeagus short and thick, apex abruptly sloping, left
paramere very large, broadly rounded at apex, right
paramere short (BA85 fig. 19).... P. monteithi Baehr
3b. Eyes as long as orbits (BA85 fig. 3c); pronotum wider,
basal angles far less acute (BA85 fig. 8); pronotum
and elytra fairly depressed; striae rather finely
punctate, punctures of intervals in 3-4 rows (BA85
fig. 5c); pilosity equal, rather depressed; aedeagus
long, narrow, and rather low, apex gently sloping,
left paramere narrow, square at apex, right paramere

rather long and narrow (BA85 fig. 15). P. brittoni

Baehr

M. Baehr

4a. Larger species (body length >5.5 mm); antenna
longer, median antennomeres well >2 x longer than
wide (BA85 fig. 4a); longer and narrower species,
fore body considerably shorter than elytra (BA85 fig.
6); aedeagus small and depressed, thickened before
apex, then gently sloping, both parameres very long

and narrow (BA85 fig. 14). P. fulvus (Castelnau)

4b. Smaller species (body length <5 mm); antenna shorter,
median antennomeres <2 x as long as wide (BA85
fig. 4b; BA95 fig. 1); shorter and wider species, fore
body almost as long as elytra (BA85 fig. 7; BA95 fig.

1) ; aedeagus far less depressed, almost equally wide,

or unknown, parameres shorter and wider (BA95
fig- 2).5

5a. Eyes smaller, orbits 1/3 x longer than eyes (BA85
fig. 3b); lateral margins of pronotum near base less
sinuate, basal angles less projecting (BA85 fig. 7);
elytra not markedly depressed on disk, 5th interval

not raised; aedeagus unknown. South Australia.

... P australis (Blackburn)

5b. Eyes larger, orbits little longer than eyes (B A95 fig. 1);
lateral margins of pronotum near base deeply sinuate,
basal angles well projecting (BA95 fig. 1); elytra
markedly depressed on disk, 5th interval distinctly
raised; aedeagus short and rather compact (BA95 fig.

2) . North Queensland. P. depressipennis Baehr

6a. Eyes at most slightly longer than orbits (BA85 fig. 3c);
pronotum and elytra rather depressed; antenna very
elongate, median antennomeres >3 x longer than wide
(BA85 fig. 4c); aedeagus elongate, depressed, apex
gently sloping, apex of left paramerc square (BA85

fig. 15). P. brittoni Baehr

6b. Eyes much longer than orbits (Figs 3, 4; BA85 figs
3d-f, h); body more convex; antenna shorter, median
antennomeres <3 x longer than wide (BA85 figs 4d-f,
h); aedeagus shorter and stouter, apex more abruptly
sloping or apical lamellae with pilose appendices (Fig.
1; BA85 figs 16-18,20).7

7a. Smaller species (body length < 5.1 mm); antenna
short, median antennomeres distinctly <2 x longer

than wide (BA85 fig. 4h).8

7b. Larger species (body length >5.35 mm); antenna
longer, median antennomeres >2 x longer than wide
(BA85 figs 4d-f).9

8a. Colour brownish to black, surface glossy; head short,
neck very distinctive; elytra shorter, <1.67 x longer
than wide and perceptibly widened towards apex;
striae very coarsely punctate, punctures of intervals
fine (BA85 fig. 5h); pilosity elongate, erect, but quite
sparse; hairs c. 2 x as long as width of an interval; apex
of aedeagus with pilose lateral appendices (BA85 fig.
20). p hirsutulus Baehr

8b. Colour reddish to light brown, surface less glossy;
head longer, neck less distinctive; elytra longer, >1.77
x longer than wide and markedly parallel-sided (Fie.
5); punctures of striae and intervals less dissimilar;
pilosity shorter and denser; apex of aedeagus without
such appendices (Fig. 1). P. parallelus sp. nov.

9a. Antenna rather elongate, median antennomeres >
2.5 x longer than wide (BA85 fig. 4d); striae finely
punctate, punctures of intervals dense, in 3-4 rows,
punctures not much smaller than those of striae (BA85
fig. 5d); usually elytra distinctly iridescent; aedeagus
very thick and broad, both parameres large, apex of
right paramere very short and wide (BA85 fig. 16)..

. P. iridescens Baehr

9b. Antenna shorter, median antennomeres c. 2 x longer
than wide, or little longer (BA85 figs 4e, fj; striae
coarsely punctate, punctures of intervals considerably
finer than those of striae (BA85 figs 5e, 0; elytra not,
or but faintly iridescent; aedeagus not so large, right
paramere longer and more acute (BA85 figs 17, 18),
or unknown.10

10a. Eyes very large, semicircular and remarkably
protruded (BA85 fig. 4f); punctures of intervals in
2-3 rows (BA85 fig. 5f); pilosity rather elongate,
unevenly erect and depressed; aedeagus fairly small
and narrow, lower surface concave, internal sac small,
right paramere very narrow and elongate (BA85

fig. 18)... .P. cyclophthalmus Baehr

10b. Eyes less semicircular and protruded (Fig. 3; BA85
fig. 4e); pilosity shorter, more regular and depressed;
either punctures of striae less coarse and punctures
of intervals at least in 3 rows, or punctures of striae
very coarse, almost as wide as the intervals, and
punctures of intervals in 2 rows; aedeagus short and
thick, widened apicad, internal sac very large, right
paramere much shorter (BA85 fig. 17), or unknown
. 11

1 la. Punctures of striae very coarse, punctures almost as
wide as one interval; punctures of intervals sparse, in
2 rows (BA85 fig. 5e); pilosity rather sparse; aedeagus
short and thick, widened apicad, internal sac very
large, right paramere much shorter (BA85 fig. 17)..

. P. punctatostriatus Baehr

lib. Punctures of striae less coarse, punctures at most
half as wide as one interval; punctures of intervals
denser, in 3 or even 4 rows; pilosity denser; aedeagus
unknown. P. gregoryensis sp. nov.

New species of Pseudaptinus from northern Australia

REMARKS

The Australian species of the genus Pseudaptinus are
remarkably similar in many aspects of body shape and
structure, except for the unique P. hirsutulus, in which
the external morphology as well as that of the aedeagus
differs considerably from all the other Australian species.
The question, why so many morphologically similar
species arc able to coexist in tropical northern Australia,
commonly at the same locality, is not solvable given the
lack of information about the ecological preferences and
habits of any species of this genus. Almost all specimens
with label data on collecting circumstances, and almost
all specimens that I have collected during several trips in
northern Australia, were sampled at light. Although it may
be supposed that all Pseudaptinus species are more or less
hygrophilous and during the daytime they probably hide
under leaf litter, or debris, or in cracks in the soil, nothing
has been ever recorded about their nutrition, courtship, or
reproduction, and the larvae are also unknown.

It would be interesting to know whether species of
Pseudaptinus can be sampled cither by systematic pitfall
trapping or by Berlese Funnel extraction of leaf litter and
debris. Both these collecting methods would probably yield
more exact information about the habitat preferences of
the species.

ACKNOWLEDGMENTS

I am grateful to the Deutsche Forschungsgemeinschaft
who supported my collecting trip and visits to certain
Australian museums under grant No. Ba 856/10-1. The
authorities of the Parks and Wildlife Commission of the
Northern Territory kindly granted me a permit to collect in
certain Northern Territory national parks. Thanks are also
due to Jan Forrest and Eric Matthews, Adelaide, and Tom
Weir, Canberra, for the loan of specimens.

REFERENCES

Baehr, M. 1984. Revision der Australischen Zuphiinae. 1. Gattung
Acrogenys Maclcay (Insecta, Coleoptera, Carabidae). Spixiana
7: 115-134.

Baehr. M. 1985a. Revision of the Australian Zuphiinae. 3. The genus
Pseudaptinus Castelnau (Insecta, Coleoptera, Carabidae).
Spixiana 8: 33-57.

Baehr, M. 1985b. Revision of the Australian Zuphiinae. 4. The
genus Parazupliium Jcannel (Insecta, Coleoptera, Carabidae).
Spixiana 8: 295-321.

Baehr, M. 1986a. Revision of the Australian Zuphiinae. 5. The genus
Zuphiunt Latreille (Insecta, Coleoptera, Carabidae). Spixiana
9: 1-23.

Baehr, M. 1986b. Revision of the Australian Zuphiinae. 6. The genus
Planetes Macleay. Supplement to the other genera (Insecta,
Coleoptera, Carabidae). Spixiana 9: 151-168.

Baehr, M. 1988. Revision of the Australian Zuphiinae 2. Colasidia
monteitlii sp. nov. from North Queensland, first record of the
tribe Leleupidiini in Australia (Insecta: Coleoptera: Carabidae).
Memoirs of the Queensland Museum 25: 135-140 (1987).

Baehr, M. 1991. On new and rare Leleupidiini from the Oriental
and Australian Regions (Coleoptera, Carabidae, Zuphiinae).
Mitteilungen der Munchner Entomologischen GesellschaJ)
81: 193-202.

Baehr. M. 1992. A new Acrogenys Macleay from Central Australia.
Supplement to the revision of the Australian Zuphiinae
(Insecta, Coleoptera, Carabidae). Spixiana 15: 75-80.

Baehr, M. 1995. A new species of Pseudaptinus Castelnau from
Australia. 2nd supplement to the ‘Revision of the Australian
Zuphiinae’ (Insecta, Coleoptera, Carabidae, Zuphiinae).
Koleoptemlogische Rundschau 65: 15-18.

Baehr, M. 2001. New species and new records of Zuphiinae
from Australia (Coleoptera: Carabidae). - Sukunahikona.
Special Publication of the Japan Coleopterological Society
1: 83-101.

Baehr, M. 2008. New species of the zuphiine genus Acrogenys
Macleay from Australia (Carabidae: Zuphiinae). Coleoptera
11 (for 2007): 113-124.

Moore, B. R 1995. Two remarkable new genera and species
of troglobiotic Carabidae (Coleoptera) from Nullarbor
Caves. Journal of the Australian Entomological Society 34:
159-161.

Accepted 9 September 2008

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 63-77

Biogeographic implications of Ascidiacea (Tunicata)
from the Wessel Islands (Arafura Sea)

PATRICIA KOTT

Honorary Associate, Queensland Museum,

PO Box 3300, South Brisbane, QLD 4101, AUSTRALIA
patricia.mather@qm.qld.gov.au

ABSTRACT

A collection of ascidians from the Wessel Islands (off north-eastern Arnhem Land) confirms the Indo-West Pacific
affinities of the Australian tropical ascidian fauna. The collection, dominated by species of the family Didemnidae
previously observed to be diverse in tropical waters, includes several of the species with obligate Prochlonm symbioses.
The collection also contains three large solitary species (Polycarpa aurita, P. papillata and Microcosmus helleri) known
to have a pan-tropical range. The geographic range and life history strategies of the majority of species in this collection
support the view that gene flow in the Indo-West Pacific and through the straits to the north of the Australian continent
that connect the two oceans is not constrained by either a short free-swimming larval life, or internal fertilisation in the
fixed, colonial organisms that dominate the tropical fauna. In colonial species at least, population maintenance may be
a more significant selective advantage than gene flow through larval dispersal.

Keywords: Indo-West Pacific, temperate, tropical, pan-tropical, Didemnidae, gene flow.

INTRODUCTION

The 35 ascidian specimens (colonies and solitary
individuals) reported on below were taken by SCUBA
from the Wessel Islands (in the Arafura Sea off north¬
eastern Arnhem Land; see Table 1). Colonial, especially
didemnid species, dominate the species list, only five of
the 25 species represented being solitary (Table 2).

The majority of the species in this collection have
been recorded previously from northern Australia and are
confirmed as common components of the benthic fauna
occurring in tropical Australian waters. The collection
contains species with a range in tropical and occasionally
temperate waters of the western Pacific from Fiji to the
eastern Australian coast (including the Great Barrier Reef),
the northwestern Australian coast and sometimes further,
occasionally to the West Indian Ocean. Only one species
(Trididemnum marmorattun) known from the western
Pacific is not yet recorded further west than the northern
coast of Australia. At present the possibly indigenous
Australian species known only from tropical seas are few
(five being present in this collection, see Table 2). Four of
the species present (including three that are solitary, viz.
Polycarpa aurita, P. papillata and Microcosmus helleri )
have a pan-tropical range.

The Ascidiacea arc sessile (fixed) organisms with larvae
free-swimming for relatively short periods and it is possible
that they could be vulnerable to isolation and spcciation.
In fact, collections (including the present one, see Table
2 below) made in shallow tropical waters, where species

diversity is high, are dominated by colonial species, which
are invariably internally fertilised, brooding their embryos
internally and with larvae free-swimming for particularly
short periods. Further, these characteristics of colonial
ascidians are particularly conspicuous in the Didemnidae,
the most speciose family in the tropical ascidian fauna, in
which larvae arc free-swimming for periods of ten minutes
or less. On the other hand, solitary species (with the
exception of some species of Polycarpa and Molgida) are
externally fertilised, embryos develop into tailed larvae in
a planktonic phase, and be more likely agents of gene flow
than the short-lived larvae of colonial species.

However, although the characteristics of the life
histories of colonial ascidian species (in particular, the
Didemnidae), tend to support an hypothesis of population
isolation resulting in their spcciation, the geographic
range of the majority of shallow water tropical ascidian
species (confirmed in the present collection) suggest that
this hypothesis of isolation is incorrect; and that gene flow
around the tropical Indian and West Pacific Oceans and
through the straits that connect these oceans, separating
the Australian continent from the islands to the north,
is not impeded by the sessile habit of the adults, nor by
internal fertilisation of eggs and brooding of embryos
within the parental organism and a short free-swimming
larval life.

Probably there are other selective advantages of a
colonial habit not directly related to gene flow, viz. patterns
and rates of growth to rapidly occupy substrata and exclude
other benthic organisms and a tendency to population

P. Kott

Table 1. Collection data for specimens reported on below (NTM, Museum and Art Gallery of the Northern Territory registration number;
NCI, National Cancer Institute voucher specimen). All locations are the Northern Territory (Australia).

NTM
reg. no.

Date

collected

Depth of

Depth

NCI No.

Latitude

Longitude

Site Description

collection

(m)

range

(m)

Habitat

Substrate

E500

0M9H2629-Y

11°47.89"S

136°29.04'E

Cotton Island, 100 m off Northern Bay,

English Company Islands.

30 Mar 04

13-16

Coral

heads

Fine sand

E501

0M9H2633-F

11°47.89"S

136°29.04‘E

Cotton Island, 100 m off Northern Bay,
English Company Islands.

30 Mar 04

13-16

Coral

heads

Fine sand

E502

0M9H2657-G

11°38.60'S

136°17.83'E

Raragala Island, large bay on SW side
of island, Wessel Islands.

30 Mar 04

17-20

Rocky

Coarse sand,
rubble

E503

0M9H2668-R

ll°33.9rs

136°22.34'E

Raragala and Guluwuru Islands
(Gugari Rip). Wessel Islands.

30 Mar 04

19-27

Channel

Rock, sand

E504

0M9H2669-S

11°38.60’S

136°I7.83'E

Raragala Island, large bay on SW side
of island, Wessel Islands.

30 Mar 04

17-20

Rocky

Coarse sand,
rubble

E505

0M9H2670-T

11°32.85'S

136°21.28‘E

Raragala Island, 700 m NE of tip,

Wessel Islands.

31 Mar 04

13-16

Muddy

bottom

Mud, rock

E506

0M9H2674-X

11°32.85'S

136°21.28'E

Raragala Island, 700 m NE of tip,

Wessel Islands.

31 Mar 04

13-16

Muddy

bottom

Mud, rock

E507

0M9H2676-Z

11°06.69'S

136°41.26'E

Trafalgar Bay Headland, Marchinbar
Island. Wessel Islands.

31 Mar 04

9-12

Rocky

reef

Rock, sand

E508

0M9H2677-A

11°06.69'S

136°41.26'E

Trafalgar Bay Headland, Marchinbar
Island. Wessel Islands.

31 Mar 04

9-12

Rocky

reef

Rock, sand

E509

0M9H2678-C

1l°06.69'S

136°41.26'E

Trafalgar Bay 1 leadland, Marchinbar
Island, Wessel Islands.

31 Mar 04

9-12

Rocky

reef

Rock, sand

E510

0M9H2687-N

1 r01.29'S

136°43.55 E

Marchinbar Island, 900 m NW of Emu
Islet. Wessel Islands.

31 Mar 04

11-17

Rocky

reef

Rock, sand

E511

0M9H2718-V

iri0.95'S

136°39.24 E

Shark Point, 1 km W of Jensen Bay,
Marchinbar Island, Wessel Islands.

02 Apr 04

17-20

Rocky

reef

Rock, silt, sand

Hopeful Bay. 340 m NW of Breakwater

10-19

Rocky

reef

Mud, rock

E512

0M9H2727-H

11°24.69'S

136°28.85'E

Point. Marchinbar Island, Wessel

Islands.

02 Apr 04

E513

OM9H2740-U

11°28.30'S

136°25.43'E

Guluwuru Island, 580 m NE of tip,
Wessel Islands.

03 Apr 04

14-15

Rocky

Rock, silt

E514

0M9H2741-V

H°41.17'S

136°02.03’E

Drysdale Island, 600 m offshore of SE
side of island, Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E515

0M9H2742-W

11°41.17S

136°02.03'E

Drysdale Island, 600 m offshore of SE
side of island. Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E516

0M9H2743-X

11°41.17'S

136°02.03'E

Drysdale Island, 600 m offshore of SE
side of island, Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E517

0M9H2744-Y

11°41.17'S

136°02.03'E

Dry sdale Island. 600 m offshore of SE
side of island, Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E518

0M9H2745-Z

11°41.17'S

136°02.03’E

Drysdale Island, 600 m offshore of SE
side of island. Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E519

0M9H2746-A

11°41.17'S

136°02.03 E

Drysdale Island, 600 m offshore of SE
side of island, Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E520

0M9H2747-C

11°41.17'S

136°02.03'E

Drysdale Island. 600 m offshore of SE
side of island, Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E521

0M9H2748-F

11°41.17’S

136°02.03'E

Drysdale Island, 600 m offshore of SE
side of island, Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E522

0M9H2749-G

11°41.17'S

136°02.03 E

Drysdale Island, 600 m offshore of SE
side of island, Wessel Islands,

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E523

0M9H2750-H

11°41.17'S

136°02.03 E

Drysdale Island. 600 m offshore of SE
side of island. Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E524

0M9H2751-I

11°41.I7’S

136°02.03'E

Drysdale Island, 600 m offshore of SE
side of island. Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E525

0M9H2753-K

11°41.19'S

I36°02.13'E

Drysdale Island. 600 m offshore of SE
side of island, Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

bommies, sand

E526

0M9H2754-L

H°41.19’S

136°02.13E

Drysdale Island, 600 m offshore of SE
side of island, Wessel Islands.

04 Apr 04

17-20

Coral

heads

Coral

Bommies, sand

E527

0M9H2768-Z

11°38.56'S

136°17.85'E

Raragala Island, 240 m NE of bay on
SW side, Wessel Islands.

05 Apr 04

11-20

Rocky

reef

Coral, sand,
rubble

E528

0M9H2782-Q

11°38.27'S

136°17.52E

Raragala Island, 600 m NE of bay on
SW side, Wessel Islands.

05 Apr 04

13-14

Sandy

bottom

Sand, mud

E529

OM9H2783-R

11°38.27'S

I36°17.52'E

Raragala Island, 600 m NE of bay on
SW side. Wessel Islands.

05 Apr 04

13-14

Sandy

bottom

Sand, mud

E530

0M9H2802-N

11°23.26'S

136°29.36'E

Marchinbar Island, 3 km N of
Breakwater Point, Wessel Islands.

02 Apr 04

15-17

Rocky

Mud, rock

E531

0M9H2803-O

11°06.69”S

136°41.26'E

Trafalgar Bay Headland, Marchinbar
Island, Wessel Islands.

31 Mar 04

9-12

Rocky

reef

Rock, sand

Ascidians from the Wessel Islands

Table 2. Species of ascidians taken from the Wessel Islands and English Company Islands, showing their previously known geographic
range and NTM registration number(s) of relevant specimens. * IWP. Indo-West Pacific; NE, northeastern Australia; NW, northwestern
Australia; WP, West Pacific.

Taxon name

Recent

reference

Previously known geographic range*

NTM
rcg. no.

No.

specs.

Suborder Aplousobranchia

Family Polycitoridae

Polycitor circes Michaelsen, 1930

Kott 2004

WP, NE, NW and northern Australia

E518

Eudistoma ehoreum Kott, 1990a

Kott 2008

NW, NE and northern Australia

E502

E513

E524

Eudistoma ovatuin (Herdman, 1886)

Kott 2004

Northern Australia

E508

E520

Eudistoma pyriforme (Herdman, 1886)

Kott 1990a

Torres Strait, NW, NE Australia

E522

Family Didemnidac

Leptoclinides braiuli Kott, 2001

Kott 2004

NE, northern Australia

E514

Leptoclinidcs cf. rigidus Kott, 2001

Kott 2005a

WP, NE, NW and northern Australia

E521

Polysyncraton oceanium Kott, 2001

Kott 2004

WP and northern Australia

E519

Didemnuin clavum Kott, 2001a

Kott 2004

WP NW Australia

E515

Didemnum molle (Herdman, 1886)

Kott 2001

IWP tropieal/temperate Australia

E527

1 +

Didemnuinpsammaiode (Sluiter, 1895)

Kott 2002

IWP tropical/temperate Australia

E510

E517

Didemnum roberli Michaelsen, 1930

Kott 2004

NW, northern Australia

E501

Didemnum viride (Herdman, 1906)

Kott 2002

IWP, NW, NE Australia

E525

Trididemnum marmoratum (Sluiter. 1909)

Kott 2004

WP, northern Australia

E516

E509

Trididemnum savignyi (Herdman, 1886)

Kott 2001

Pan-tropical, NW, NE Australia

E526

Trididemnum sibogae (Hartmeyer 1910)

Kott 2007

IWP tropical / temperate Australia

E529

Lissoclinum badium F. and C. Monnot, 1996

Kott 2004

WP, NW, NE Australia

E523

Lissoclinum multijidum (Sluiter, 1909)

Kott 2004

IWP, northern and southern Australia

E503

Suborder Phlebobranchia

Family Ascidiidae

Pliallusia arabica Savigny, 1816

Kott 1985

IWP, NW, NE Australia

E530

Family Perophoridae

Perophora modificata Kott, 1985

Kott 2004

WP, NE Australia, Ashmore Reef

E507

Suborder Stolidobranchia

Family Stvelidae (Subfamily Styelinae)

Polycarpa aurita (Sluiter 1890)

Kott 2008

Pan-tropical, NW, NE Australia

E594

Polycarpa longiformis Tokioka, 1952

Kott 1985

NW, NE Australia, Japan

E500

Polycarpa papillata (Sluiter, 1885)

Kott 2006

Pan-tropical, tropical/temperate

E505

Australia

E512

Family Styelidae (Subfamily Polyzoinae)

Stolonica australis Michaelsen, 1927

Kott 1985

Temperate Australia

E528

Symplegma brakenhielmi (Michaelsen, 1904)

Kott 2004

IWP and temperate Australia

E531

Family Pyuridae

Microcosmus helleri Herdman. 1882

Kott 1985

Pan-tropical, tropical/temperate

E511

Australia

E506

maintenance by concentration of recruits (resulting from
reduction in the larval exposure to dispersal, albeit retaining
larvae that are free-swimming for minimum periods for
site selection). The maintenance of stable populations may
enhance opportunities for fertilisation (both internal and
external) that transcend the advantages of increasing gene
flow through a longer larval life; and form links in chains
of recruitment around the margins of the continents and

between the islands and reefs of the western Pacific and
Indian Oceans.

Unfortunately, at this stage there is little data available
on the patterns of life histories, population dynamics and
selective advantages reflected in the dominant groups of
the diverse ascidian fauna in the tropical Indo-West Pacific
(see Kott 1985, 1990,2001; glossary entries for fertilisation
and gene flow).

P. Kott

BIOGEOGRAPHIC NOTES

Generally the species of ascidians represented in this
collection are those that would be expected in the north of
Australia. They can be divided into the following groups:

1. Species with a wide pan-tropical range. Microcosmus
helleri and Polycarpa papillata are known to extend
also around Australia into temperate waters, but
Trididemnum savignyi and Polycarpa aurita do not
extend further south than the tropics.

2. Species known from the tropical West Pacific
(including north-eastern Australia) and the Indian
Ocean. For some species in this group, the only Indian
Ocean record is offnorth-westem Australia ( Polycitor
circes, Leptoclinides cf. rigidus, Didemnum clavum,
Lissoclinum badium and Pewphora modificata), but
others ( Didemnum molle, Didemnum viride, Phallusia
arabica, Didemnum psammatode, Trididemnum
sibogae and Symplegma brackenhielmi) extend to the
West Indian Ocean and the last two species listed also
extend from the tropics into temperate waters around
the southern coast of Australia. Lissoclinum multifidum
is recorded from the western Pacific, the Indian Ocean
and northern Australia as well as southern Australia,
but has not been recorded from north-eastern or north¬
western Australia and it is possible that more than one
species is involved.

3. Only one species is known only from the western Pacific
including northern eastern Australia (Trididemnum
marmoratum ).

4. Species, possibly indigenous, known from the north
of Australia, including the north-western and / or
north-eastern coasts ( Eudistoma eboreum, E. ovatum,
E. pyriforme, Leptoclinides brandi, Polysyncraton
oceanium, Didemnum roberti, Polycarpa longiformis)
have not been recorded outside Australian waters.

5. Stolonica australis is the only species not previously
reported outside temperate Australian waters and
the present new record from the Wessel Islands is an
extension of its known geographic range.

TAXONOMIC NOTES

The specimens in this collection were photographed
in situ by their respective collectors. Unfortunately, the
photographed specimen (Figs 1-3) is not necessarily
the same specimen as the one examined and reported on
here.

Polycitor circes Michaelsen, 1930

(Fig.l A)

Polycitor circes Michaelsen, 1930: 495; Millar 1975:
205, part, specimens from Marongas (20.iii.14); Monniot
1988:207; F. and C. Monniot 1996: 184; F. and C. Monniot
2001: 249; Kott 1990a: 169 and synonymy; Kott 2002:
26. Not Millar 1975: 205 part, specimen from Marongas

(19.iii. 14) and those from other locations (see Remarks,
below).

Distribution. Previously recorded (see Kott 1990a):
Western Australia (north-western Australia, Shark Bay,
Cockbum Sound); Queensland (Martha Ridgeway Reel);
Northern Territory (Darwin), Papua New Guinea, New
Caledonia, Indonesia, Philippines. New record: Northern
Territory (Northern Territory (Wessel Islands, NTM
E518).

Description. The newly recorded colony is a firm
gelatinous translucent vertical column, expanded slightly at
the top where the test is slightly softer and more translucent
than the firm test that forms the stalk. The surface of the
colony is smooth and even.

The zooids extend from the upper surface of the head
where each zooid opens to the exterior by two separate
6-lobed apertures. They extend parallel to one another
toward the base of the stalk. Zooids arc very long, with a
long oesophageal neck. The abdomen continues past the
pole of the gut loop and terminates in a vascular stolon.
Longitudinal muscles extend along the length of the zooid
and terminate in a short horn-like protrusion on each side
of the anterior part of the posterior abdominal stolon
(posterior to the pole of the gut loop). About 25 rows
of stigmata are in the large thorax. The stomach, at the
posterior end of the abdomen, has about 20 longitudinal
folds. The gonads are in the gut loop and consist of a
few large eggs and many small testis follicles scattered
through and around the pole of the gut loop. Large
embryos arc present moving up the oviduct as it extends
up the oesophageal neck toward the atrial cavity. Larval
adhesive organs are triradially arranged at the anterior
end of the trunk.

Remarks. The present colony resembles many assigned
to Diazona futtgia (see F. and C. Monniot 2001, Fig.
123E). However it is readily distinguished by the absence
of internal longitudinal vessels in the branchial sac, the
conspicuous folds in the stomach (rather than the internal
striations of Diazona) and by the embryos moving up
through the abdomen (fertilisation having taken place in
the base of the oviduct).

Colonies from Deo Roa (Philippines) with small flat-
topped lobes branching off basal stolons assigned to this
species by Millar (1975) are probably specimens of Clavelitut
arafurensis Tokioka, 1952. They arc quite different from
the massive robust vertical lobes of the type specimens of
the present species (Michaelsen 1930) and others described
by Kott (1990a) from Western Australia. Another specimen
Millar (1975) described from Deo Roa with smooth margins
around the apertures appears to be a Pycnoclavella sp. Of
the other specimens Millar (1975) assigned to this species,
one from Marongas (20.iii.14, Millar 1975) appears to
be properly assigned, although the other specimen from
that location (19.iii.l3) without stomach folds may be a
specimen of Polycitor translucidus.

Ascidians from the Wessel Islands

Fig 1. A, Polycitor circes ; B, Eudistoma eboreum ; C, Eudistoma ovatunv, D, Leptoclinides cf. rigidus; E, Didemnum clavum', F, Didemnum
psammatode. Photos: A, P. Colin; B, D-F, D. de Maria; C, M. Browne.

P. Kott

The specimens assigned to Polycitor giganteus by
F. and C. Monniot (2001) and Kott (2005b) from New
Caledonia and the Solomon Islands, respectively, lacking
stomach folds and having larval adhesive organs in a
median vertical line, probably are also specimens of
Polycitor translucidus. Polycitor giganteus Herdman,
1899 has zooids of similar proportions to the present
species but is known mainly from temperate Australian
waters and has only four stomach folds (sec Kott
1990a).

Although not described, specimens from Papua New
Guinea appear to have been accurately assigned to Polycitor
circes by F. and C. Monniot (2001). Specimens from New
Caledonia and Indonesia (F. Monniot 1988; F. and C.
Monniot 1996) also appear to be members of the present
species.

Eudistoma eboreum Kott, 1990
(Fig. IB)

Eudistoma eboreum Kott, 1990a: 205; Kott 2002: 27;
Kott 2008: 1118.

Distribution. Previously recorded (see Kott 2008):
Western Australia (Kalbarri); Northern Territory (Darwin);
Queensland (Lizard Island). New records'. Northern
Territory (Wessel Islands, NTM E502, E513, E524).

Description. The newly recorded colonies are purple-
brown, translucent, firm gelatinous slabs with evenly
distributed groups of about four to six zooids arranged in
a circle with their atrial apertures opening to the surface
in the centre of the circle and their branchial openings
around the circumference. In the living colonies there
is some variation in the extent to which these systems
are depressed into the surface, although in preserved
specimens the surface is more or less even. Also
photographs of two of the newly recorded specimens
show the colonies to have been a whitish colour and pale,
but the other colony (NTM E513) is distinctly grey-blue.
Collector’s notes describe one of these colonies (NTM
E524) to have been brown-black internally. However,
in preservative all three specimens are grey with diffuse
brown pigment in the test of the upper half of the colony.
Zooids have the characteristic layer of transverse muscles
in the thoracic body wall that overlies strong longitudinal
bands extending the length of the long zooids. Atrial
apertures are on a long siphon. Three rows of stigmata
are in the branchial sac, which is separated from the
smooth-walled stomach at the posterior end of the zooid
by a long oesophageal neck. A small vascular stolon is
at the posterior end of the body.

Remarks. Although the colour of living specimens of
this species is variable, their colour in preservative is more
or less the same. The robust zooids in circular rudimentary
systems in the firm grey translucent slabs of test are
characteristic of this species. It is known from relatively few
specimens across the northern tropical coast of Australia.

Eudistoma ovation Herdman, 1886
(Fig. 1C)

Eudistoma ovation Herdman, 1886: 246; Kott 2004: 42
and synonymy. Not Kott 1990a: 222 and synonymy (see
E. pyriforme below).

Distribution. Previously recorded (see Kott 2004):
Northern Territory (Torres Strait, Gulf of Carpentaria,
Bynoe Harbour). New records : Northern Territory (Wessel
Islands, NTM E508: 3 specimens, E520: 1 specimen).

Description. Colonies are vertical to conical or rounded
lobes to 2 cm diameter and 3 cm high, sessile or with a
thick short stumpy basal stalk. Muddy-looking plant cells,
faeces pellets and sand arc embedded in the soft test around
the long robust zooids that extend from the upper surface
toward the base of the colony. The zooids are arranged in
circles with the long atrial siphons opening in the centre
of the circle and the branchial openings in an outer circle.
Zooids are muscular with an outer layer of transverse
muscles and inner longitudinal bands along the length of
the zooids, which extend from the upper surface toward the
base of the colony. As is usual in this genus, the oesophageal
neck is very long. In this species, the gut forms a spiral in
the posterior end of the loop.

Remarks. The spiral in the gut loop and the inclusions
of various particles in the test are characteristic of this
species. The former character distinguishes the species
from the otherwise similar Eudistoma amplum (Sluiter,
1909). Polycitor multiperforatus Sluiter, 1909 also is a
similar species, although it appears to have more stigmata
per row (30) and the circular arrangement of zooids was
not detected in the syntypes (see Kott 1990a).

Eudistoma pyriforme (Herdman, 1886)

Psammaplidium pyriforme Herdman, 1886: 419.

Eudistoma pyriforme. — Kott 1990a: 226 and
synonymy.

Eudistoma ovation. -— Kott 1990a: 222.

Distribution. Previously recorded (see Kott 1990a):
Western Australia (Onslow); Queensland (Bundaberg);
Torres Strait. New record'. Northern Territory (Wessel
Islands, NTM E522).

Description. The colony is a flat slab with the margins
raised around the flattened upper surface. Fine sand is on the
upper surface, interrupted where the zooids open separately
to the exterior. Sand also is present throughout the colony.
Some excrescences or root-like projections are around the
sides of the colony enmeshed with sand and other particles.
Zooids are contracted and the abdomen is contracted to
about the same length as the thorax. Three rows of stigmata
were detected in the branchial sac but other details of the
zooids were not observed. Two small larvae arc in the thorax
of the newly recorded specimen.

Remarks. The present colony is not unlike those
previously reported for this species. Although some sand
and possibly symbionts are embedded in the test, it lacks
the upright lobes, the conspicuous circular systems and

Ascidians from the Wessel Islands

the spiral at the distal end of the gut loop of E. ovatum, a
species that often has been confused with it.

Leptoclinides brandi Kott, 2001

Leptoclinides brandi Kott, 2001a: 40. — Kott 2004:
2468.

Distribution. Previously recorded (see Kott 2004):
Queensland (Great Barrier Reef); Northern Territory
(Darwin and Bynoe Harbour). New record : Northern
Territory (Wessel Islands, NTM E514).

Description. The tough, fleshy-looking colony has a
smooth surface raised into mounds and ridges. A layer
of large, stellate spicules (to 0.08 mm diameter) with
9-13 moderately long and pointed conical rays in optical
transverse section are in the surface of the colony but the
remainder of the translucent test has only sparse spicules.
In life, the colony is greyish white. The arrangement of the
common cloacal systems is obscured by compression of the
colony. Zooids are very contracted, although the branchial
siphon is long with a bulbous expansion halfway along it.
The atrial siphon is long and oriented posteriorly. Stigmata
are relatively short in these contracted zooids but they
appear to be pointed at each end. About six male follicles
are in a circle and the vas deferens coils five times.

Remarks. The newly recorded colony resembles those
previously described, although its condition obscures
some of its characters, especially the fusiform stigmata.
The stigmata and their distribution arc characteristic of
the species and these together with the testis follicles and
coils of the vas deferens distinguish the species from those
in the dubius group, which resemble the present colony in
some respects.

Leptoclinides cf. rigidus Kott, 2001
(Fig. ID)

Leptoclinides rigidus Kott, 2001 a: 77; Kott 2005a: 2425
and synonymy.

Distribution. Previously recorded (see Kott 2005a):
Western Australia (Ashmore Reef, Montebello Islands);
Queensland (Great Barrier Reef); Northern Territory
(Darwin, Wessel Islands); Papua New Guinea. New record:
Northern Territory (Wessel Islands, NTM E521).

Description. Colonies arc gelatinous sheets overgrowing
the substrate. The small zooids are arranged along each side
of canals that surround zooid-free circular areas about 3
mm diameter, each containing a patch of spicules beneath
a superficial bladder cell layer. Spicules are sparse in
the remainder of the test. Large sessile common cloacal
apertures are at the junctions of some of the common cloacal
canals, which all contain some faecal pellets. The spicules
are to 0.05 mm diameter with 7-9 conical rays in optical
transverse section. Some of the ray tips are relatively blunt.
In life, the colonies arc a reddish brown. Little of the zooid
structure can be determined. As in all species of this genus a
posteriorly orientated atrial siphon is present and a retractor
muscle was not detected

Remarks. Superficially the specimen resembles the
temperate Leptoclinides rigidus , which has been recorded
previously from this location. However specimens
previously assigned to the species are reported to have been
blue in life. The present specimen resembles Didemnum
viride in life, but it could be an undescribed species of the
genus Leptoclinides. Leptoclinides volvus Kott, 1975 has
similar common cloacal systems, but it is a stalked colony
and is also a temperate species.

Polysyncraton oceanium Kott, 2001

Polysyncraton oceanium Kott, 2001a: 115.

Distribution. Previously recorded (see Kott 2001a):
Queensland (Great Barrier Reef); Fiji. New record:
Northern Territory (Wessel Islands, NTM E519).

Description. The colony is an investing sheet with
firm translucent test. A thin layer of evenly spaced but not
crowded spicules is in the surface test beneath a superficial
layer of bladder cells. A sparse layer of spicules also is on
the base of the colony but there are no spicules in the test
between the surface and basal layers. Spicules are stellate,
to 0.035 mm diameter, with 11-15 short pointed rays that
break up readily. A common cloacal cavity was not detected
in this specimen, although the zooids sometimes were seen
to be arranged in double rows, presumably along each side
of canals. Five coils of the vas deferens were detected, but
the gonads were not seen, although some large eggs are
present in the basal test.

Remarks. The form and distribution of the spicules
resemble those of Didemnum caesium Sluiter, 1909,
Polysyncraton rica Kott, 2001. P otuetue C. & F. Monniot,

1987 and P. scobinum Kott, 2001, but they are smaller. The
specimen generally conforms to those previously assigned
to Polysyncraton oceanium.

Didemnum davitm Kott, 2001

(Fig. IE)

Didemnum clavum Kott, 2001a: 163; Kott 2004: 53.

Distribution. Previously recorded (see Kott 2004):
Western Australia (NW Australia to Port Hedland);
Northern Territory (Darwin); Indonesia. New record:
Northern Territory (Wessel Islands, NTM E515).

Description. The colony is the characteristic branched
structure with red pigment particles in the surface and
spicules crowded throughout. The spicules are small,
stellate, with occasional giant spicules, with 4-6 long
pointed rays, scattered amongst them

Remarks. Although other tropical Didemnum species
are a similar red-pink colour, the present species is readily
identified by the form of the colonies and the occasional
giant spicules with relatively few rays scattered amongst
the smaller stellate spicules.

P. Kott

Didenutum /nolle (Hcrdman, 1886)

Diplosomoides molle Herdman, 1886: 310.

Didemnum molle. — Kott 2001: 208 and synonymy.

Distribution. Previously recorded (see Kott 2001):
NW, NE and northern Australia to Cockburn Sound and
one record from Esperance. Western Pacific, the Indian
Ocean (to Mauritius, Madagascar), Vietnam. New record:
Northern Territory (Wessel Islands, NTM E527).

The newly recorded location is well within the vast
geographic range of this shallow reefal species, one of
the most frequently recorded of the didemnid-Proc/i/oron
symbioses.

Description. The species is readily identified, forming
vase-shaped colonies with a central test core separated
from the outer zooid-bearing layer of the colony by a
vast common cloacal cavity lined by green symbiotic
cells, which can be seen through the large terminal
common cloacal aperture. The species characteristically
secretes vast quantities of mucus in which the green
symbiotic cells are liberated from the colony when it
is disturbed.

Remarks. The species is discussed in Kott (2001).

Didemnum psammatode (Sluiter, 1895)

(Fig. IF)

Leptoclinum psamathodes Sluiter, 1895: 171.

Didemnum psammatode. — Kott 2002: 38.

Distribution. Previously reported (see Kott 2002): a
wide recorded range in the Indian and Pacific tropical/
temperate oceans between Sri Lanka, the Red Sea to Fiji
and extending to southern Australia and the South China
Sea. New records'. Northern Territory (Wessel Islands, NTM
E510, E517)

Description. The newly recorded specimens are as
previously described, with mud-coloured faecal pellets
crowded in the test and minute spicules around each zooid
opening.

Didemnum roberti Michaelsen, 1930
(Fig. 2A)

Didemnum roberti Michaelsen 1930: 516; Kott 2004:
58 and synonymy.

Distribution. Previously recorded (see Kott 2004):
Western Australia (north west to Cockburn Sound);
Northern Territory (Gulf of Carpentaria, Darwin, Torres
Strait). New record : Northern Territory (English Company
Islands, NTM E501).

The species is commonly recorded from the
northwestern coast of the continent and from Darwin,
the Gull of Carpentaria and Torres Strait, but it is not yet
reported from northeastern Australia or from locations
in the Western Pacific. At this stage, its records suggest
that it is one of the few indigenous Australian tropical
species.

Description. The colony is robust with terminal large
circular common cloacal apertures on the rounded surface

swellings, branches and ridges. The surface is always
smooth and even. Spicules are in a relatively sparse layer
at the surface and are more sparsely distributed and often
patchy internally. They also are in a sparse but even layer
lining the extensive common cloacal cavity that separates
the central test core from the surface zooid-bearing layer
of the colony. Zooids arc in clumps, each attached by a
short basal connective that traverses the common cloacal
cavity from the centre of the under surface of each clump.
Each zooid has a fine retractor muscle. The atrial aperture
is large, open and sessile and the vas deferens coils eight
times around the undivided testis follicle. Larvae, with
four pairs of lateral ampullae along each side of the
antero-median adhesive organs, are embedded in the
basal or central test.

Remarks. This species has robust but variable
colonies with a similarly variable colour pattern. It
resembles many species of Leptoclinides in its smooth
outer surface and well-developed common cloacal
systems with a large terminal common cloacal aperture
and an extensive posterior abdominal cavity separating
a central test core from an outer zooid-bearing layer.
Assignation of the species to the genus Didemnum is
confirmed by characteristics of zooids, such as the large
open sessile atrial apertures; the fine tapering retractor
muscle; and the undivided testis surrounded by the coiled
vas deferens.

Didemnum viride (Herdman, 1906)

(Fig. 2B)

Leptoclinum viride Herdman, 1906: 340.

Didemnum viride. — Kott 2001a: 9.

Distribution. Previously reported (see Kott 2001a):
Western Australia (Montebello Islands); Queensland (Great
Barrier Reef); Papua New Guinea, New Caledonia, French
Polynesia, Indian Ocean (Sri Lanka). New record: Northern
Territory (Wessel Islands, NTM E525).

Description. In preservative, the newly reported colony
is a thin cream-coloured sheet with the surface marked
by a mosaic of small elevated areas separated by narrow
depressions over the common cloacal canals that are lined
on each side by zooids. In life, the colony is brownish red
in the depressed areas over the common cloacal canals.
Spicules, to 0.05 mm diameter, are crowded in the basal
half of the colony but are less crowded in the zooid layer.
They have 7- 11 conical rays with more or less rounded tips
in optical transverse section.

Remarks. The specimen generally resembles Didemnum
poecilomorpha F. & C. Monniot, 1996 although it lacks the
two different types of spicules of the latter species. In the
present collection the colony resembles that of Leptoclinides
cf. rigidus (see above), but the oval elevations that form
the mosaic on the surface of the colony are smaller in the
present species.

Ascidians from the Wessel Islands

Fig 2. A, Didemnum roberti; B, Didemnum viride; C, Trididemnum marmoratum ; D, Trididemnum sibogae; E, Lissoclinum badiunv
F, Lissoclinum multifidum. Photos: A,B,D,E, D. de Maria; C,F, P. Colin.

P. Kott

Trididemttum marmoratum (Sluiter, 1909)

(Fig. 2C)

Leptoclinum marmoratum Sluiter, 1909: 84.

Trididemnum marmoratum. — Kott 2002: 38; Kott
2004: 61.

Distribution. Previously reported (see Kott 2004):
Northern Territory (Gulf of Carpentaria, Darwin);
Indonesia. New records'. Northern Territory (Wessel Islands,
NTM E509, E516).

Description. One of the newly reported colonies (E5 1 6)
has relatively uniform rounded elevations about 1.0 cm
diameter at the base and up to 0.5 cm high, each with a
terminal common cloacal aperture. The other specimen
is an extensive encrusting sheet. Stellate spicules, to 0.09
mm diameter, with 9-13 conspicuously pointed or chisel¬
shaped rays in optical transverse section are in a thin layer
beneath the superficial bladder cell layer but are absent from
the remainder of the colony. Primary posterior abdominal
common cloacal spaces beneath the surface layer of zooid-
bearing test are traversed by clumps of zooids and surround
a central test core in each of the conical elevations. Zooids
are robust, with a posteriorly orientated atrial siphon
opening into the posterior abdominal common cloacal
cavity. Sometimes they have black squamous epithelium
on the thorax and abdomen, although this was not always
detected. Occasionally there is also a black endostylar
pigment cap. A short retractor muscle is present from the
posterior end of the thorax. A large spherical male follicle
is surrounded by at least eight coils of the vas deferens.
Larvae with a narrow waist behind a corona of eight club-
shaped ectodermal ampullae with bifid tips on each side of
the three antero-median adhesive organs are in the central
test core.

Remarks. The specimens resemble closely those
previously assigned to this species by Kott (2002).

Trididemnum savignyi (Herdman, 1886)

Didemnum savignyi Herdman, 1886: 281.

Trididemnum savignyi. — Kott 2001a: 64.

Distribution. Previously reported (see Kott 2001a):
Western Australia (Nares Rock); Queensland (Great Barrier
Reef); Northern Territory (Darwin); tropical Atlantic. New
record: Northern Territory (Wessel Islands, NTM E526).

Description. The newly recorded colony is an extensive
thin sheet growing over a spiral worm tube, although the
in situ photograph of this specimen is difficult to reconcile
with its appearance in preservative. A sparse layer of large
spicules, to 0.09 mm diameter, with 9-11 conical pointed
rays in optical transverse section is beneath a superficial
bladder cell layer. Spicules also line the oesophageal
common cloacal canals but are absent from the remainder
ol the colony. Faecal pellets arc embedded in the basal
layer of test. Zooids have large funnel-shaped atrial siphons
from about halfway down the dorsal surface of the thorax.
Sometimes abdomina have dark squamous epithelium.

Remarks. The newly recorded specimen resembles
previously reported specimens of this species, although the
dark pigment previously thought to be characteristic is not
present. This is presumably associated with intraspecific
variability in this widely distributed species.

Trididemnum sibogae (Hartmeyer, 1910)

(Fig. 2D)

Didemnum sibogae Hartmeyer, 1910: 261.

Trididemnum sibogae. — Kott 2001a: 283 and
synonymy; Kott 2007: 1203 and synonymy.

Distribution. Previously recorded (see Kott 2007):
Western Australia (Cape Jaubert); South Australia (Cape
Jaffa); Victoria (Western Port); Tasmania (Port Davey);
New South Wales (Port Hacking, Port Jackson, Arrawarra);
Queensland (Great Barrier Reef and mainland locations
between Fraser Island and Princess Charlotte Bay); Northern
Territory (Darwin, Gulf of Carpentaria). Indonesia, New
Caledonia, Indian Ocean (Gulf of Manaar). New record'.
Northern Territory (Wessel Islands, NTM E529).

The records indicate this species to be common in
temperate as well as tropical waters around Australia.

Description. The newly recorded colony is a fist-sized
three-dimensional reticulum (sec Kott 2001a, Fig. I32A)
with thin branches overgrowing and fusing with one another
to form the sponge-like consistency of the colony. An in
situ photograph shows the colony to be creamish-ycllow.
The colony has a superficial bladder cell layer and spicules
are scattered relatively sparsely through the colony. An
extensive posterior abdominal common cloacal cavity
separates the zooid-bcaring surface layer of test to the
central test in each of the branches. Spicules are relatively
small, to 0.06 mm diameter, stellate, with 7-11 conical,
sharply pointed rays in optical transverse section. Zooids
are small, with a short posteriorly oriented atrial siphon.

Remarks. The large, 0.016 mm diameter, spicules
reported to occur occasionally in this species were not
detected in the present specimen, although otherwise its
characters conform to those previously reported.

Lissoclinunt badium F. and C. Monniot, 1996

(Fig. 2E)

Lissoclinum badium F. and C. Monniot, 1996: 170;
Kott 2004: 69.

Distribution. Previously recorded (see Kott 2004):
Western Australia (Bonaparte Archipelago); Queensland
(Great Barrier Reef); Northern Territory (Darwin); Coral
Sea, Timor Sea, Palau Is. New record : Northern Territory
(Wessel Islands, NTM E523).

Description. The characteristic soft flexible specimen,
dark brown internally, has large common cloacal apertures
on surface ridges that are prominent on the living inflated
colony. Specimens of this species arc readily identified.
Brown pigment cells are mixed with the especially small
spicules in the surface of the colony and brown coloured
zooids can be seen crossing the extensive thoracic common

Ascidians from the Wessel Islands

cloacal cavity. The basal half of the colony is opaque
with crowded spicules but the abdomina of the zooids are
embedded in the floor of the common cloacal cavity where
the spicules arc less crowded.

Lissodinum miiltifidum (Sluiter, 1909)

(Fig. 2F)

Leptodinum multifidum Sluiter, 1909: 311.

Lissoclinum multifidum. — Kott 2004: 65 and
synonymy.

Distribution. Previously recorded (see Kott 2004):
South Australia (Flinders Island, etc); Tasmania (Forestier
Peninsula); Northern Territory (Darwin); Indonesia, Gulf
of Thailand, Mauritius. New record: Northern Territory
(Wessel Islands, NTM E503).

Description. T he colony is said to have been brown in
life although in the in situ photograph it looks red. It encrusts
rubble, some of which is mixed with the basal or central
test mass. This is surrounded by an extensive common
cloacal cavity crossed by test commissures that attach it
to the zooid-bearing layer of test. Large common cloacal
apertures are on surface elevations. Small sparse spicules,
to 0.025 mm diameter, with a range of spicule rays from
conical to rounded or rod-like are scattered through the test
together with morula cells and possibly some plant cells as
described previously for this species.

Remarks. Although Kott (2001a) had thought this
species to be aspicular, it docs in fact have minute spicules
scattered through the test.

Phallusia arabica Savigny, 1816
(Fig. 3A)

Phallusia arabica Savigny, 1816: 164; Kott 1985: 61;
Kott 2001b: 61 and synonymy.

Distribution. Previously recorded (see Kott 1985):
Queensland (from the Capricorn Group, southern Great
Barrier Reef, to Trinity Bay), Arafura Sea, Philippines,
Sri Lanka, Red Sea, Gulf of Suez. New record: Northern
Territory (Wessel Islands, NTM E530).

Description. The newly repotted specimen has a robust,
slightly curved body attached to the substrate about halfway
down the convex ventral surface with a short conical atrial
protuberance from the concavity about halfway down
the dorsum. Anteriorly the body narrows to the terminal
branchial aperture. The test is smooth, translucent and
relatively firm and thick, and both atrial and branchial
siphons appear to be linn and inflexible. However, internally
the siphons are long and muscular, and the body lies loosely
in the test and does not adhere to it. A mesh of longitudinal
and transverse muscles is over the right side of the body and
on the left a similar mesh is present but only anterior to the
gut loop. Branchial tentacles are short and darkly pigmented
in the preserved specimen. The long dorsal ganglion is at
the base of the atrial siphon. A small U-shaped slit is on the
dorsal tubercle at the anterior end of the dorsal lamina but a
peritubercular-V is shallow. Very small secondary openings

of the neural duct open into the peribranchial cavity along
the left side of the dorsal lamina, which has conspicuous
ribs along the left side of the membrane. The gut, on the
left side of the posterior half of the body, is filled with mud.
The rim of the anal opening is divided into about 20 short
rounded scallops.

Remarks. This robust, widely ranging species is readily
distinguished from Phallusia obesa (Herdman, 1880), P.
julinea Sluiter, 1919 and P millari Kott, 1985, all commonly
occurring species of this genus in the Indo-West Pacific,
by the smooth surface of the present species, the lack of
chromatophores in the test, the presence of the dorsal
ganglion at the base of the atrial aperture rather than halfway
between the atrial opening and the dorsal tubercle and the
lack of the sandy holdfast characteristic of P. millari.

Perophora modificata Kott, 1985
(Fig. 3B)

Perophora modificata Kott, 1985: 104; Kott 2004: 40
and synonymy.

Distribution. Previously reported (see Kott 2004):
Western Australia (Ashmore Reef); Queensland (Great
Barrier Reef), Coral Sea Plateau; Palau Islands, Philippines.
New record: Northern Territory (Wessel Islands, NTM
E507).

Description. The newly recorded specimen is a small
clump of rounded yellow zooids each on a long stalk
attached to a basal mat of stolons. The branchial sac has
four rows of stigmata. The zooids in this and all other
respects conforms to previous accounts of this readily
identified species.

Polycarpa aurita (Sluiter, 1890)

(Fig. 3C)

Styela aurita Sluiter, 1890: 338.

Polycarpa aurita. — Kott 1985: 152 and synonymy;
Kott 2008: 1199.

Distribution. Previously reported (see Kott 1985):
Western Australia (Cape Jaubert, Shark Bay to Cockbum
Sound); New South Wales (Port Jackson); Queensland
(Moreton Bay to Lizard Island); Northern Territory (Gulf
of Carpentaria); Indonesia, New Caledonia, Philippines;
Atlantic Ocean (Venezuela, Gulf of Mexico, Caribbean).
New record: Northern Territory (Wessel Islands, NTM
E504).

The species has not yet been reported from the eastern
Pacific, the Indian or the eastern Atlantic Oceans, but
otherwise it is pan-tropical.

Description. The newly recorded specimen is robust
with a hard leathery test. It is large and appears to be
senescent, the gonads being degenerate, only traces being
detected embedded deeply in the body wall. The species
is identified principally by its large gut, large elliptical
stomach, the strong ligament joining the two limbs of the
primary loop, the long crowded vertical endocarps between
the limbs of the gut loop and, on the body wall, the small

P. Kott

rounded scallops lining the rim of the anus and the deep
conspicuous slit on the dorsal tubercle.

Remarks. The specimen is cryptic, the leathery wrinkled
test being covered with epibionts and in the field it can be
confused with other stolidobranch species in this collection,
such as Microcosmus helleri, although it does not appear to
form aggregates as many of these species do.

Polycarpa longiformis Tokioka, 1952
(Fig. 3D)

Polycarpa longiformis Tokioka, 1952: 119; Kott 1985:
170 and synonymy.

Distribution. Previously reported (see Kott 1985):
Western Australia (Port Hedland); Queensland (Martha
Ridgeway Reef, Lizard Island, Cape Weymouth, Raine
Island, Murray Island); Arafura Sea, Japan. New record :
Northern Territory (English Company Islands, NTM
E500).

Description. The newly reported specimen is large
(about 4 cm long), almost cylindrical, grey and rubbery-
looking with a smooth surface. The test, though thin, is
firm and the muscular body wall adheres closely to it.
The apertures are both anterior on conical protuberances
diverging from one another. The lobes of the aperture are
completely obscured by the firm gelatinous test. Posteriorly
the body narrows to a holdfast attached to rocks and other
nibble. The branchial folds are narrow and widely separated
with 5-7 internal longitudinal vessels in the interspace
and the meshes are long. The gut forms a short loop in
the posterior end of the body and the rectum continues
anteriorly to the base of the atrial siphon. The anal border
has long, rounded lobes. Loosely attached long polycarps
in an uneven row down each side of the body are obscured
by the long vertical teardrop-shaped cndocarps crowded
on the body wall.

Remarks. In the field, the gelatinous smooth conical
protuberances with their terminal apertures projecting from
crevices can be mistaken for the very similar structures in
Phallusia species (P. julinea, P. arabica and P. obesa) and
in specimens of Polycarpa papillata. The close relationship
with P. papillata is discussed by Kott (1985).

Polycarpa papillata Sluiter, 1885
(Fig. 3E)

Styela (Polycarpa) papillata Sluiter, 1885: 192.

Polycarpa papillata. — Kott 1985: 184 and synonymy;
Kott 2006: 218.

Distribution. Previously reported (see Kott 1985):
Western Ausralia (Dampier Archipelago to Cockburn
Sound); South Australia (St Vincent Gulf); Victoria
(Portland); New South Wales (Port Jackson); Queensland
(Morcton Bay to Bathurst Island); Northern Territory
(Gulf of Carpentaria, Darwin); Arafura Sea, Indonesia,
Palau Islands, Philippines, Marianas Islands, Sri Lanka,
Madagascar. New records : Northern Territory (Wessel
Islands, NTM E505, E512).

Description. Long, tough specimens, pink in preservative
with faint pink longitudinal stripes. The outer test is smooth
and even without ridges or creases. The thin but muscular
body wall is closely adherent to the test. The body narrows
to the anterior terminal branchial opening which is turned to
the right and slightly downwards. The atrial aperture is on
a conical protrusion projecting out at an angle to the body
about one-third of the distance down the dorsal surface.
Branchial folds are long and narrow and widely separated
from one another. The body wall has crowded upright leaf¬
like cndocarps, flattened and sometimes indented around
their rounded margins and constricted at their bases where
they join the body wall. They are especially crowded around
the gut loop in the posterior part of the body. The anus has
16 long and slightly frilly anal lobes around the margin.
Long polycarps in about three irregular rows are embedded
in the body wall or attached to it by a long ligament at their
proximal ends, but the dorsal end of each polycarp with its
terminal gonoducts is free of the body wall and directed
toward the atrial opening.

Remarks. The species is common in the tropical Indo-
West Pacific and although it resembles P longiformis, it
is readily identified by its red colour and longitudinal red
stripes. It often has ejected its gut when collected although
in the newly recorded specimens the body organs are all
present.

Stolonica australis Michaelscn, 1927

Stolonica australis Michaelscn, 1927: 202; Kott 2003:

1642 and synonymy.

Distribution. Previously recorded (sec Kott 2003):
Western Australia (Albany); South Australia (Great
Australian Bight, St Vincent Gulf, Investigator Strait,
Kangaroo Island, Yorke Peninsula); Tasmania (Bruny
Island. d’Entrecasteaux Channel); Victoria (Bass Strait,
Anglesea, Portland, Western Port); New South Wales (Port
Jackson). New record: Northern Territory (Wessel Islands,
NTM E528).

Description. Zooids are vertical with a terminal
branchial aperture and the atrial aperture antero-dorsal, each
on a slight conical protuberance. Zooids are joined to one
another and to basal stolons by short lateral commissures
to form tight aggregations that are brownish pink in life.
The test is thin but rigid. Two branchial folds and a possible
third ventral rudimentary fold are on each side of the body.
A typical branchial formula is 9DL4(12) 5(9) 9E. A total
of about 40 internal longitudinal vessels are on each side.
The stigmata are in about 20 rows. The dorsal tubercle is
a longitudinal slit. About 25 longitudinal folds are in the
stomach wall, of which six terminate on each side of the
suture line. Only a short caecum projects from the pyloric
end of the suture line into the gut loop. Small cndocarps are
scattered on the body wall and in the gut loop. Small male
follicles are scattered in an arc around the postero-ventral
curve of the right side of the body.

Ascidians from the Wessel Islands

Fig 3. A, Phallusia arabica; B, Perophora modificata ; C, Polycarpa aurita ; D, Polycarpa longiformis : E, Polycarpapapillata: F, Microcosmus
helleri. Photos: A-C, D. dc Maria, D,E, P. Colin; F, B. Alvarez de Glasby.

P. Kott

Remarks. The present species has formerly been
regarded as having a range limited to temperate waters, but
the newly recorded material suggests that the range may be
more extensive and include tropical waters. It does have
some similarities with Stolonica alula Kott, 1985 (recorded
from the south-western part of Western Australia), which
has a similar zooid with a long vertical slit on the dorsal
tubercle and similar distribution of gonads around the
postero-ventral curve of the body, but three folds on each
side of the body, a longer gastric caecum and a gastric spur
that extends the stomach into the gut loop.

Symplegma brakenhielmi (Michaelsen, 1904)

Diandrocarpa brakenhielmi Michaelsen, 1904: 50.

Symplegma brakenhielmi. — Kott 2004: 71.

Distribution. Previously recorded (sec Kott 2004):
Western Australia (Cape Preston to Cockburn Sound);
Queensland (Moreton Bay to Martha Ridgeway Reef).
Indonesia, Noumea, Palau Islands, Fiji, Thailand, Hong
Kong, China, Indian Ocean including Sri Lanka. New
record'. Northern Territory (Wessel Islands, NTM E531).

Description. The colony forms a large robust sheet
over the substrate. Zooids are dorso-ventrally flattened and
completely embedded in the thin test forming a mosaic like
pattern seen from the upper surface. Apertures are sessile
on the upper (dorsal) surface of each zooid. Characteristic
of the genus are the four longitudinal branchial vessels on
each side of the branchial sac. The gut forms a tight double
loop on the left side of the body and the stomach is large,
but relatively short and wide with a conspicuous curved
caecum in the pole of the primary gut loop. The two testis
follicles, one anterior and one posterior to the small ovarian
sac in the middle of the body wall, are lobed. Larvae are in
the peribranchial cavity of the newly recorded specimen.

The species and its relationships are discussed in Kott
(2004).

Microcosmus Itelleri Herdman, 1882
(Fig. 3F)

Microcosmus helleri Herdman, 1882: 131; Kott 1985:
349.

Distribution. Previously recorded (see Kott 1985):
Western Australia (Port Hcdland, Cockburn Sound); South
Australia (St Vincent Gulf); Indonesia. Java Sea, Singapore,
Sri Lanka, West Indian Ocean, West Indies. New records'.
Northern Territory (Wessel Islands, NTM E506, E511).

The species is pan-tropical but also extends into
temperate waters around the southern coast of Australia.

Description. The newly recorded specimens, up to
about 9 cm long, are tough, hard and leathery, with wrinkled
test and some scattered epibionts. The branchial aperture is
terminal and turned slightly ventrally. Internally the atrial
siphon extends from near the base of the branchial siphon
and turns posteriorly, lying at almost 180 degrees to it,
opening in the sessile external atrial aperture about halfway
down the dorsal surface of the body. Hard, pointed tubercles

of the test arc around both external openings. Four hard,
rounded and slightly spoon-shaped valves are around the
base of the branchial siphon and each is folded back against
the lining of the siphon from its base just anterior to the
branchial tentacles. Siphonal spines were not detected in
these specimens. The dorsal tubercle has the double spiral
slit characteristic of several species in this genus. Six wide
branchial folds are on each side of the body. The gut forms
a narrow loop around the ventral margin of the left side and
the gonads, divided into three large blocks embedded in the
body wall pass from inside the pole of the gut loop, cross
its descending limb and continue alongside anterior to the
gut loop to the atrial aperture.

Remarks. The course of the left gonad, crossing from
inside the pole of the loop and extending close to the rectum,
occurs also in M. exasperatus Heller, 1878 and M. squamiger
Michaelson, 1927. However, this large, tough pan-tropical
species, which extends into temperate waters around the
southern Australian coast, is distinguished by the lack of
siphonal spines, the presence of four rather cartilaginous
spoon-like valves in the base of the branchial siphon and
the six branchial folds on each side of the body.

ACKNOWLEDGMENTS

The specimens were collected by Dr Belinda Alvarez
de Glasby of the Museum and Art Gallery of the Northern
Territory and her party (M. Browne, H. Nguyen, D. de
Maria and R Colin) under the ‘Collection and Taxonomy
of Shallow Water Marine Organisms’ program for the
US National Cancer Institute (Contract N02-CM-27003).
Access to the area was kindly granted by Mr Terry
Yumbulul, who also provided the team with advice on
collection sites. I am grateful to Dr Alvarez de Glasby for
making the collection and the in situ photographs of the
specimens available to me.

I am grateful also to the Trustees and the CEO of the
Queensland Museum (Dr Ian Galloway) for providing
the accommodation and other infrastructure that make it
possible for me to pursue this work.

REFERENCES

Hartmeyer, R. 1909-1911. Ascidien (continuation of work by
Sccligcr). Pp. 1281-1772. In: Bronn, H.G. (ed.) Klassen und
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Accepted 10 November 2008

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 79-86

Juvenile bull sharks Carcharhinus leucas (Valenciennes, 1839)
in northern Australian rivers

DEAN C. THORBURN AND ANDREW J. ROWLAND

Centre for Fish and Fisheries Research, Murdoch University,

South St, Murdoch, Western Australia 6150, AUSTRALIA
Corresponding author: dthorhurn@aapt.net.au

ABSTRACT

Juvenile bull sharks, Carcharhinus leucas , ranging in total length from 687 mm to 1365 mm, were captured in riverine
waters throughout the Northern Territory and Kimberley region. Western Australia. Carcharhinus leucas was captured
in highest abundance in low salinity riverine waters (<5 ppt), however, none were captured in offshore marine waters.
The rivers of northern Australia act as nurseries for juvenile C. leucas within which individuals remain for approximately
four years. Assuming annuli present on vertebrae were laid down annually and a birth size of approximately 700 mm
TL, growth estimates suggest that C. leucas attains 863 mm TL in its first year, 993 mm TL in its second year, 1118
mm TL in its third year and 1239 mm TL in its fourth year. Stomach content analysis indicated a broad diet with the
consumption of large amounts of teleost fishes within those systems, as well as numerous other prey types including
terrestrial mammals, aquatic insects and other elasmobranchs.

Keywords: bull shark, nursery area, Carcharhinus , river, Kimberley, Northern Territory.

INTRODUCTION

The occurrence of the bull shark Carcharhinus leucas
in Australian rivers is well documented as it is the only
carcharhinid known to penetrate well into freshwater
systems for extended periods of time (Whitley 1943;
Chubb et al. 1979; Merrick and Schmida 1984; Last and
Stevens 1994; Allen et al. 2002). However, records of the
occurrence of C. leucas in offshore waters of Australia are
rare, a disparity that may be attributed to the confusion of
this species with other whalers and in particular with the
very similar pigeye shark ( Carcharhinus amboinensis)
(Cliff and Dudley 1991; Last and Stevens 1994).

Although biological data for Australian populations
of C. leucas are scarce, various aspects of the life history
have been studied elsewhere including Lake Nicaragua
(Thorson et al. 1966; Thorson and Lacy 1982), the Florida
Lagoons (Snclson et at. 1984), Gulf of Mexico (Branstetter
and Stiles 1987) and South Africa (Cliff and Dudley 1991;
Wintner et al. 2002). These studies indicate that C. leucas
can remain in freshwater systems for extended periods and
often spends a significant part of its juvenile life in upper
estuarine and freshwater reaches (Thorson 1972; Snelson
et al. 1984; Cliff and Dudley 1991). Breeding is thought
to occur in offshore marine waters, with females entering
estuarine and inshore waters to give birth (Montoya and
Thorson 1982; Bishop et al. 2001). Length at birth estimates
range from 55 to 90 cm total length (Thorson and Lacy
1982; Snelson 1984; Branstetter and Stiles 1987; Cliff and
Dudley 1991; Wintner et al. 2002).

Despite the marine affinities of C. leucas, this species'
utilisation of upper riverine reaches has often led to it being
included in studies of freshwater elasmobranchs. This was
the case during two previous ichthyological surveys that
aimed to capture elasmobranchs in fresh waters of northern
Australia (Taniuchi et al. 1991; Ishihara et al. 1991). As a
result of these studies and other surveys of riverine waters
C. leucas has been shown to occupy numerous systems
in tropical and warm temperate regions of Australia
extending south to Sydney (New South Wales) and Perth
(Western Australia) (Chubb et al. 1979; Last and Stevens
1994; Bishop et al. 2001; Allen et al. 2002; Thorbum et al.
2003).

In light of the paucity of knowledge on elasmobranchs
occurring in estuarine and freshwaters of northern Australia,
ichthyological surveys were conducted in the Northern
Territory' and Kimberley region of Western Australia in 2002
(Thorbum et al. 2003) and additionally in the Fitzroy River
(Western Australia) throughout 2003 and 2004 (Thorbum et
al. 2004). During these surveys, C. leucas was captured in
higher abundance than any other elasmobranch species, and
provided an opportunity to: determine the distribution and
any broad-scale habitat associations of C. leucas in the rivers
of the Northern Territory and Kimberley region, Western
Australia; and describe the biology, age composition and
diet of C. leucas occurring within these rivers. Furthermore,
this study aimed to test the hypothesis that juveniles of
the species utilise rivers of northern Australia as nursery
grounds, by satisfying the criteria defined by Heupel et al.
(2007) that, “(1) sharks are more commonly encountered

D. C. Thorburn and A. J. Rowland

in the area than other areas; (2) sharks have a tendency to
remain or return for extended periods; and (3) the area or
habitat is repeatedly used across years”.

MATERIALS AND METHODS

Study sites, environmental variables and habitat. One

hundred and eighty-six marine, estuarine and freshwater
sites were sampled between the Robinson River (NT)
and the Fitzroy River (WA) from June 2002 to July 2004
(Fig. 1). The salinity (ppt), temperature (°C), and an estimate
of water clarity using a secchi disc (cm) were recorded at
each sample site. The depth (m) and tidal movement were
also recorded. Notes on the immediate habitat, including
predominant sediment type, density of aquatic vegetation
types and detritus, riparian vegetation and snag density,
were also made.

Sample collection, measurements and field
dissection. Sampling was primarily conducted with sinking
monofilament gill nets (including 20 m panels of 5, 7.5, 10,

15 and 20 cm stretched mesh) that were set perpendicular to
the river bank. All gill net mesh sizes were utilised during
daylight hours, however, only the larger meshes were
used at night to minimise by-catch. Baited long-lines and
handlines were also used. Long-lines were a maximum of
40 m long and contained up to 20 5/0 tuna circle hooks.
Each hook was connected to the 500 lb nylon mainline via
0.5 m of 200 lb wire trace. Locally available fish was used
as bait. Long-lines were set during the day and night.

Fig. 1. The sites at which Carcharhinus leucas was captured in the Northern Territory and in the Kimberley region of Western Australia.
Included are records from the Museum and Art Gallery of the Northern Territory (NTM) and the Western Australian Museum (WAM).

Dissections were conducted in the field as soon as
practicable after capture to minimise tissue breakdown
and further digestion of the gut contents. The total length
(TL, mm), sex and weight (W, g) were recorded for each
specimen captured. For comparison with other studies
which reported in precaudal length (PCL) only, the
conversion TL (cm) = (PCL + 9.16) / 0.81 (Branstetter and
Stiles 1987) was used.

The stomach and a minimum of six vertebrae from
below the first dorsal fin were removed from 51 female and
49 male C. leucas. Stomach fullness and contributions were
either determined soon after capture or the stomachs were
preserved whole in 100% ethanol for later examination.
Vertebral samples were stored in 100% ethanol or kept on
ice until they could be frozen.

Length-weight, length-frequency and age. A
likelihood ratio test (Cerrato 1990) was used to determine
if the length-weight relationship was significantly different
between the sexes. As no difference appeared to exist the
sexes were pooled and the SPSS statistical package used
to determine the line of best fit for the relationship. A
length-frequency histogram was generated to aid in the
identification of size specific cohorts.

Samples of vertebrae were defrosted and placed in 5%
sodium hypochlorite solution until free of tissue. The centra
were then rinsed thoroughly in water and allowed to dry for
several hours. A minimum of two centra from each sample
was embedded in resin and a 0.3 mm longitudinal section
cut with an Isomet low speed rotary saw. Sections were
then mounted on a slide with DePex mounting medium
and observed under a dissection microscope with reflected

Juvenile bull sharks ( Carcharhinus leucas ) in northern Australian rivers

light. Counts of the number of growth rings or annuli
commencing after the birth mark (identified by a change
of angle on the outer edge of the corpus calcerium) on the
facia of whole and sectioned vertebrae were then made
for each individual. The number of annuli was compared
to TL and the mean size of individuals with respect to the
number of annuli calculated. The lack of successive annual
samples of vertebrae precluded validation that annuli were
laid down annually during the current study. Marginal
increment analysis of vertebrae from specimens from the
Gulf of Mexico (Branstetter and Stiles 1987) and ‘mark-
recapture’ analysis of captive South African specimens
(Wintner et at. 2002) confirmed the annual formation of
annuli. These findings, in conjunction with fact that length
at age and growth rate data collected during the current
study (see Results) were comparable to those collected by
Branstetter and Stiles (1987) and Wintner et at. (2002),
suggest that annuli may similarly be laid down annually in
Australian specimens.

A von BertalanfTy growth curve was fitted to the length
of C. leucas at their estimated age of capture, however,
this method required the provision of the individual’s birth
dates. This estimation was subsequently based on the stage
of healing of the umbilical scars present on a number of
the individuals captured. Healing rates were assumed to be
comparable to that of Carcharhinus cautus where umbilical
scars were observed to be completely healed within three
months of birth (Dr William White, CSIRO, Hobart, pers.
comm). The birth date of an individual possessing an open
umbilical scar was estimated to have been at the beginning
of the month of capture. An individual possessing a closed
umbilical scar was assumed to have been born at the
beginning of the month prior to the date of capture, and
an individual possessing a near faded umbilical scar was
assumed to have been born two months prior to capture.
Other individuals collected from the same river as those
possessing umbilical scars, or from other rivers in close
proximity, were subsequently assigned the same birth
months.

Based on the individuals and times captured, those
from (and near to) the Roper River (NT) were assumed
to be born I June, those from the Daly River (NT) on 1
August, those from the Ord River (WA) on 1 November,
and those from the Fitzroy River (WA) on 1 February. A
line of regression was fitted with the use of SigmaPlot 8.0
and the von Bertalanffy growth parameters generated by
the equation L = L r (1 - e' A ’ ( ''" l) ), where L t is the length at
age t, L : is the mean asymptotic length, K is the growth
coefficient and t (j is the hypothetical age at which the
estimated length is zero.

Maturation. Maturation in male C. leucas was
determined on the basis of clasper calcification. Individuals
were considered immature when claspers were small
and uncalcified, maturing if claspers were extending and
becoming semi-calcified and mature when claspers were
fully calcified and sperm was present. Maturity in females

was assessed on observations of the ovaries and uteri.
Individuals possessing undeveloped ovaries and thin,
flaccid uteri were considered immature, maturing when the
uterus began to enlarge and ovaries contained differentiated
ova, and mature when the ovaries contain yolked ova and
an enlarged uterus (Conrath 2004).

Diet. Upon removal of the stomach, an estimate of
fullness on a scale of 0 to 10 (zero representing an empty
gut and 10 being fully distended) was made. Some smaller
prey items required identification under a dissection
microscope. An estimation of the percentage contribution
of each food item was made for each individual, and
the frequency of occurrence (%F) and mean percentage
volumetric contribution (%V) calculated (Hynes 1950;
Hyslop 1980).

The graphical method of Costello (1990) was also
used to display the broad functional prey categories found
to constitute the diet of C. leucas ; %F is plotted against
relative quantity (in this case %V). This method takes
into account the percentage occurrence, and subsequently
provides insight into population wide food habits (Cortes
1997). Unidentified items were excluded from analysis and
the values of all other dietary 1 categories of that individual
adjusted upward to sum to 100% (Pusey et al. 2000). Similar
prey categories were combined to reflect a functional prey
group.

RESULTS

Capture locations, sex ratio, length ranges and
habitat. A total of 111 C. leucas were captured from 41
of the 186 sites sampled (Fig. 1). Fifty seven were female
and 54 were male (sex ratio of 1.06 female : 1 male) which
ranged in length from 691 to 1382 mmTLand 687 to 1365
mm TL, respectively (Fig. 2). Carcharhinus leucas was
only captured within rivers (i.e. upstream of the mouth).
The sites of capture ranged in salinity from 0 to 41.1 ppt
(Fig. 3), however, 86% of individuals (all of which were

Fig. 2. Length-frequency histogram for female and male Carcharhinus
leucas collected during the study.

D. C. Thorbum and A. J. Rowland

immature) were captured in waters of less than 5 ppt (32%
were captured in 0 ppt). Carcharhinus leucas was caught
in waters ranging in temperature from 20.8 to 32.5 °C that
were between 0.7 and 21 nr deep and which varied greatly in
clarity (5-400 cm secchi depth). The species was generally
captured in open waters over sandy substrates that supported
low macrophyte and algal growth.

Length-weight relationship, age and growth,
maturity and diet. Weights of C. leucas ranged from
2360 to 20 500 g in females and from 1740 to 18 000 g
in males (Fig. 4). The relationship between TL and W can
be described by the formula W = 6165.14 - (21.741TL) +
(0.0228TL 2 ).

Observations of the annuli present on the vertebrae
of 100 individuals suggested the presence of five year
classes (0+ to 4+) (Fig. 5). The average TL of individuals
possessing zero annuli on vertebrae was 787 mm,
individuals possessing one annuli on vertebrae averaged
859 mm, individuals possessing two annulus on vertebrae

o —

Salinity (ppt)

Fig. 3. The number of Carcharhinus leucas captured versus sample
site salinity (the number of sites sampled are given in parenthesis).

W = 6165.14 - (21.741TL) + (0.0228TL 2 )

Total length (mm)

Fig. 4. The relationship between total length (TL) (mm) and weight
(W) (g) of Carcharhinus leucas captured in northern Australia.

1600

1400

| 1200

JZ
Ul
g

S 1000
£

800

600

0(31) 1(30) 2(13) 3 (22) 4(4)

Number of annuli

Fig. 5. Number of annuli observed on vertebrae versus the total length
(TL) of Carcharhinus leucas. The number of individuals possessing
0 to 4 annuli are given in parentheses.

averaged 1010 mm, individuals possessing three annuli on
vertebrae averaged 1 1 68 mm and individuals possessing
four annuli on vertebrae averaged 1339 mm.

The von Bcrtalanffy growth equation generated
parameter estimates of L a = 4662 mm, K = 0.0347 and t„
= -4.88 years (Fig. 6). While the L r is above the known
maximum size of-3400 mm TL (Last and Stevens 1994),
both the / 0 and L m are comparable lo those published for this
species elsewhere. From this equation it is estimated that
C. leucas attains 863 mm TL in its first year, 993 mm TL
in its second year, 1118 mm TL in its third year and 1239
mm TL in its fourth year.

All 49 male C. leucas captured possessed small non-
calcified claspers indicating immaturity. Similarly all
51 females dissected were found to be immature by the
possession of undeveloped ovaries and thin, flaccid uteri.

Seventy-six of the 100 stomachs inspected contained
prey. Teleost fishes were the major constituents of the
diet of C. leucas (Table 1, Fig. 7). Where identification
of teleost prey to species was possible, the lesser salmon
catfish Ariopsisgraeffei, bony bream Nematalosa erebi and
scaly croaker Nibea squamosa were the major components

Age (years)

Fig. 6. von Bcrtalanffy growth curve for Carcharhinus leucas
collected during the study.

Juvenile bull sharks ( Carcharhinus leucas) in northern Australian rivers

2. Crocodylus johnstoni

Ariid catfish#

3. Terrestrial mammal

4. Insect

5. Macrobrachium rosenbergii

Other teleost •

■

• Nematalosa erehi

•

#Plqnt matter

0 10 20 30 40 50 60

Occurrence (%F)

Fig. 7. The important broad functional prey categories of Carcharhinus
leucas.

Tabic 1. Percentage volumetric contribution (%V) and percentage
occurrence (%F) of the different food items found in the stomachs of
Carcharhinus leucas.

Prey Item

% V

Vegetation

Terrestrial vegetation

1.04

13.16

Aquatic macrophyte

0.22

3.95

Macrocrustacea

Macrobrachium rosenbergii

1.40

7.89

Insect

Orthoptera

0.03

1.32

Coleoptera

0.63

5.26

Teleost

Ariopsis graeffei

26.71

32.89

Ariopsis midgleyi

2.41

1.32

Hemiarius diodes

3.56

2.63

Plicofollis argypleuron

1.27

1.32

Other ariid sp.

10.11

17.11

Harpadon translucens

1.22

1.32

Hemiramphidae (garfish)

0.20

1.32

Lates calcarifer

3.21

5.26

Liza tade

2.08

2.63

Nematalosa erebi

15.14

19.74

Nibea squamosa

9.44

7.89

Polydactylus macrochir

2.86

1.32

Rhinomugil nasutus

0.95

2.63

Sclerophages jardinii

1.26

1.32

Unidentified teleost parts

6.77

21.05

Elasmobranchii

Pristis microdon

2.54

1.32

Mammal

Pig

1.59

2.63

Other

0.98

2.63

Reptile

Chelonidae

0.54

1.32

Crocodylus johnstoni

2.96

2.63

Unidentified

0.89

3.95

of the diet. The removal of the ‘unidentified’ prey category
and consolidation into broad (functional) prey categories
further reflected the dominance of teleost species in the
diet of C. leucas.

DISCUSSION

Distribution and habitat. Carcharhinus leucas
was frequently captured in fresh waters several hundred
kilometres from the coast. The notion that northern
Australian rivers act as nurseries for juveniles of the species
is strongly supported, and satisfies the definition criteria
defined by Heupel et al. (2007) in that: high abundances
were encountered upstream of the rivers mouths while none
were captured from marine waters; all specimens were
small and immature and appeared to remain in the river
until approximately three or four years of age (assuming
annuli were laid down annually); and juveniles of less than
one year were captured from the Fitzroy River in each of
the three consecutive years of sampling.

Saline estuarine waters have generally been considered
to provide important nursery habitats of C. leucas more so
than freshwaters (Simpfendorfer 2005). During the current
study however, the highest abundances of individuals were
captured in low salinity waters near the tidal limit and in
non-tidal freshwaters, suggesting that in Australia low
salinity waters are more important as nursery habitats than
higher salinity waters associated with the lower estuary.

The use of upstream waters by juvenile C. leucas is
likely attributed to the high abundance of food resources
available and lack of predators including large sharks and
estuarine crocodiles Crocodylus porosus (Simpfendorfer
and Milward 1993; Morgan et al. 2004). Upstream
migration of the species was demonstrated on several
occasions where the highest abundances of individuals
were encountered immediately below a barrier such as an
exposed rock bar, road crossing or barrage. In a number of
cases, these barriers also represent the upper limit of tidal
influence and may partially explain the high abundance of
this species in those reaches.

Size and age structure of C. leucas in northern
Australian rivers. The structure of riverine populations
in northern Australia appears similar to those observed
elsewhere including North America where C. leucas resides
in inland waters of the Florida Lagoons for a similar period
of time (Snelson et al. 1984). Studies by Wintncr et al.
(2002) and Cliff and Dudley (1991) suggested that C. leucas
in South Africa are born slightly larger than elsewhere,
including the coastal lagoons of Florida, Nicaragua and
the Gulf of Mexico, i.e. 800 to 900 mm TL cf. 600 and
750 mm TL, respectively (Snelson et al. 1984; Thorson
and Lacy 1966; Branstettcr and Stiles 1987). During the
current study, open umbilical scars were observed on
individuals between 687 and 749 mm TL. The size at

D. C. Thorbum and A. J. Rowland

birth of Australian C. leucas was therefore found to be
comparable to individuals of the latter locations.

Umbilical scars were observed in individuals captured
from the Roper River (NT) in June, Daly River (NT) in
August, Ord River in November (WA) and from the Fitzroy
River (WA) in February. Bishop el al. (2001) suggests that C.
leucas in Australia are born during summer which coincides
with the early portion of the tropical wet season. This
apparent westward progression of birthing across northern
Australia may therefore be a response to the westward onset
of the wet season (considered to be between October and
April in the NT and between January and March in WA)
(Bureau of Meteorology, Commonwealth of Australia
2004). High river discharges associated with the wet season
subsequently provide the opportunity for juvenile C. leucas
to penetrate far upstream before the contraction of water
levels during the tropical the dry season.

Various studies have indicated that the annual growth
rate of C. leucas is greatest in the first few years of life
(Caillouet 1969; Thorson and Lacy 1982; Branstctter and
Stiles 1987; Wintner et al. 2002). Assuming that annuli were
laid down annually, length at age estimates generated during
this study of863 mm TL at one year of age and 1239 mm TL
at four years of age are consistent with those observed from
the Gulf of Mexico (Branstetterand Stiles 1987). However,
length at age estimates of northern Australian specimens

appear lower than those from South Africa where a one
year old C. leucas was estimated to be 1040 mm TL and a
four year old estimated to be 1406 mm TL (Wintner et al.
2002). The current study also estimated that annual growth
in the first four years may be approximately 1 35 mm yr 1 .
This annual growth estimate is marginally greater than that
observed in South Africa (an average of 110 mm yr 1 in the
first three years) (Wintner et al. 2002), however, it is less
than the growth rate observed in Nicaragua (an average of
160-180 mm yr' in the first two years) (Thorson and Lacy
1982) and the Gulf of Mexico (an average of 150 - 200 mm
yr 1 for the first 5 years) (Branstetter and Stiles 1987).

Occurrences of sharks greater than two metres in length
were reported from near Fitzroy Crossing on the Fitzroy
River (Western Australia) over 300 km upstream of the
mouth. Furthermore, the senior author observed a C. leucas
of approximately two metres TL swimming immediately
below Camballin Barrage approximately 150 km from the
mouth of the Fitzroy River. Based on the von Bertalanffy
growth equation and length weight relationship lor younger
C. leucas generated during this study, these specimens
would be 11 years old and weigh 54 kg. Size of maturity
data collected from elsewhere has indicated that C. leucas
attains sexual maturity at between 1600 and 2200 mm TL
in males and between 1800 and 2250 mm TL in females
(Thorson et al. 1966; Bass 1977; Branstetter and Stiles

rat ' -ml

m*L

Fig. 8. Stomach contents and potential prey of C. leucas, including (a) freshwater sawfish Pristis microdon, (b) other C. leucas,
(c) freshwater crocodile Crocodylus johnstoni and (d) feral pig.

Juvenile bull sharks ( Carcharhinus leucas) in northern Australian rivers

1987; Cliff and Dudley 1991). Based on the aforementioned
observations and the length at maturity estimates, the
presence of mature C. leucas in northern Australian rivers
can not be dismissed.

Feeding behaviour and predation. Inspection of the
stomach contents of C. leucas indicated the consumption
of a wide variety of prey types and an opportunistic often
indiscriminate hunting pattern. While a vast majority of the
diet consisted of bony fishes (in particular ariid catfishes),
unusual prey including mammals (pig), other elasmobranchs
(.Pristis microdon) and portions of freshwater crocodiles
(Crocodylus johnstoni) were observed (Fig. 8). Vegetation
(aquatic and terrestrial) was also encountered in a high
proportion of the stomachs inspected, further reflecting
the random nature of prey selection. The authors were also
witness to cannibalism by the species whereby C. leucas
captured in gill nets were attacked by others. Several
C. leucas were also observed leaping approximately 2.5 m
clear of the water to take birds perched on overhanging
branches in the Fitzroy River, Wester Australia.

Anecdotal accounts of aggressive feeding described by
traditional owners of the Fitzroy River may also suggest the
presence of cooperative hunting behaviours. Of particular
interest was an account of an attack on a feral pig that had
entered the waters edge to drink. As explained, several small
C. leucas attacked the legs of the pig and proceeded to pull
it into deeper water. The pig eventually drowned and was
consumed. An additional report of an attack by C. leucas on
a freshwater crocodile was also collected. A single C. leucas
was observed to rapidly disable the crocodile by biting off
one of its front legs. Numerous other C. leucas subsequently
appeared and proceeded to bite the remaining limbs before
attacking the body.

ACKNOWLEDGMENTS

Sincere thanks must go to Dr David Morgan, Dr Howard
Gill, Dr Peter Last, Dr John Stevens, Dr Helen Larson, Dr
Barry Russell, Steven Gregg and family, Stirling Peverell,
Dr William White, Michael Taylor and Matt Pember, the
Natural Heritage Trust, Murdoch University, Department
of Fisheries Government of Western Australia, Northern
Territory Fisheries, Museum and Art Gallery of the Northern
Territory, Kimberley Land Council and Department of
Conservation and Land Management.

REFERENCES

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freshwater fishes of Australia. CSIRO/Western Australian
Museum: Perth.

Bass, A. J. 1977. Long-term recoveries of tagged sharks. Copeia 3:
574-575.

Bishop, K. A., Allen, S. A., Pollard, D. A. and Cook, M. G. 2001.
Ecological Studies on the Freshwater Fishes of the Alligator
Rivers Region, Northern Territory: Autecology. Supervising
Scientist Report 145, Supervising Scientist: Darwin.

Branstctter, S. and Stiles, R. 1987. Age and growth of the bull shark,
Carcharhinus leucas, from the northern Gulf of Mexico.
Environmental Biology of Fishes 20: 169-181.

Caillouet, C. W. 1969. Weight, length and sex ratio of immature bull
sharks, Carcharhinus leucas , from Vermillion Bay, Louisiana.
Copeia 1: 196-197.

Cerrato, R. M. 1990. Interpretable statistical tests for growth
comparisons using parameters in the von Bertalanffy equation.
Canadian Journal of Fisheries and Aquatic Sciences 47:
1416-1426.

Chubb, C. F., Hutchins, J. B., Lcnanton, R. C. J. and Potter I. C. 1979.
An annotated checklist of the fishes of the Swan-Avon river
system. Western Australia. Records of the Western Australian
Museum 8: 1-55.

Cliff, G. and Dudley, S. F. J. 1991. Sharks caught in the protective gill
nets off Natal, South Africa. 4. The bull shark Carcharhinus
leucas Valenciennes. South African Journal of Marine Science
10:253-270.

Conrath, C.L. 2004. Reproductive biology. In: Musick, J. A. and
Benlll, R. (Eds) Elasmobranch Fisheries Management
Techniques, APEC Fisheries Working Group. Pp. 133-164.

Cortes, E. 1997. A critical review of methods of studying fish feeding
based on stomach contents: application to elasmobranch
fishes. Canadian Journal of Fisheries and Aquatic Science
54: 726-738.

Costello, M. J. 1990. Predator feeding strategy and prey importance:
a new graphical analysis. Journal of Fish Biology 36:
261-263.

Ebert. D. A. 2003. California Natural History Guides. Sharks. Rays
and Chimaeras of California. University of California Press,
Berkeley.

Heupel, M. R., Carlson, J. K. and Simpfcndorfer, C. A. 2007.
Shark nursery areas: concepts, definition, characterization
and assumptions. Marine Ecology Progress Series 337:
287-297.

Hynes, 11. B. N. 1950. The food of sticklebacks with a review of the
methods used in studies of food in fishes. Journal of Animal
Ecology? 19: 36-58.

Hyslop, E. J. 1980. Stomach content analysis - A review of methods
and their application. Journal of Fish Biology 17: 411-429.

Ishihara, H., Taniuchi, T„ Sano, M. and Last, P. R. 1991. Record of
Pristis elavata Gannan from the Pentecost River, Western
Australia, with brief notes on its osmoregulation, and
comments on the syslematics of the Pristidae. University
Museum, University of Tokyo, Nature and Culture 3: 43-53.

Last, P. R and Stevens, J. D. 1994. Sharks and Rays of Australia.
CSIRO Division of Fisheries, CSIRO, Australia.

Merrick, J. R. and Schmida, G. E. 1984. Australian Freshwater
Fishes. Biology and Management. Griffin Press: Nctley, South
Australia.

Montoya, R. V. and Thorson, T. B. 1982. The bull shark (Carcharhinus
leucas) and largetooth sawfish (Pristis perotteti) in Lake
Bayano, a tropical man-made impoundment in Panama.
Environmental Biology of Fishes 7: 341-347.

Morgan, D. L., Allen, M. G., Bedford, P. and Horstman, M. 2004. Fish
fauna of the Fitzroy River in the Kimberley region of Western
Australia - including the Bunuba, Gooniyandi, Ngarinyin,
Nyikina and Walmajarri Aboriginal names. Records of the
Western Australian Museum 22: 147-161.

Pusey, B. J., Read, M. G. and Arthington, A. H. 2000. The dry-season
diet of freshwater fishes in monsoonal tropical rivers of Cape
York Peninsula, Australia. Ecology of Freshwater Fishes 9:
177-190.

D. C. Thorbum and A. J. Rowland

Sintpfendorfer, C. A., Freitas, G. G„ Wiley, T. R. and Heupel, M. R.
2005. Distribution and habitat partitioning of immature bull
sharks (Carcharhinus leucas) in a southwest Florida estuary.
Estuaries 28: 78-85.

Simpfendorfcr, C. A. and Mil ward. N. E. 1993. Utilisation of a tropical
nursery area by sharks of the families Carcharhinidac and
Sphymidae. Environmental Biology of Fishes 37: 337-345.

Snelson, F. F., Mulligan, T. J. and Williams, S. E. 1984. Food
habitats, occurrence, and population structure of the bull shark,
Carcharhinus lcucas, in Florida coastal lagoons. Bulletin of
Marine Science 34: 71-80.

Taniuchi, T., Shimizu, M., Sano, M., Baba, O. and Last, R R. 1991.
Description of freshwater elasmobranchs collected from three
rivers in northern Australia. University Museum, University of
Tokyo, Nature and Culture 3: 11-26.

Thorbum, D. C., Peverell, S., Stevens, J. D., Last, R R. and Rowland,
A. J. 2003. Status of Freshwater and Estuarine Elasmobranchs
in Northern Australia. Report to the Natural Heritage Trust.

Thorbum, D. C„ Morgan, D. L., Rowland, A. J. and Gill, H. S. 2004.
Elasmobranchs in the Fitzroy River, Western Australia. Report
to the Natural Heritage Trust.

Thorson, T, B. 1972. The status of the bull shark, Carcharhinus leucas,
in the Amazon River. Copeia 3: 601-605.

Thorson, T. B. and Lacy, E. J. 1982. Age, growth and longevity of
Carcharhinus leucas estimated from tagging and vertebral rings.
Copeia 1: 110-116.

Thorson. T. B., Watson, D. E. and Cowan, C. M. 1966. The status of
the freshwater shark of Lake Nicaragua. Copeia 3: 385-402.
Whitley, G. R 1943. Ichthyological descriptions and notes. The
Proceedings of the Linnean Societ V of New South Wales 68:
114-144.

Wintner, S. P.. Dudley, S. F. .1, Kistnasamy, N. and Everett, B. 2002.
Age and growth estimates for the Zambezi shark, Carcharhinus
leucas, from the east coast of South Africa. Marine and
Freshwater Research 53: 557-566.

Accepted 15 April 2008

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 87-146

Review of the Dinematichthyini (Teleostei: Bythitidae) of the Indo-west Pacific.
Part IV. Dinematichthys and two new genera with descriptions of nine new species

PETER RASK M0LLER 1 AND WERNER SCHWARZHANS 2

1 Natural History Museum of Denmark, Zoological Museum, University of Copenhagen,
Universitetsparken 15, DK-2100 Copenhagen 0, DENMARK
pclrmoller@snm.ku.dk

2 Ahrensburger Weg 103 D, 22359 Hamburg, GERMANY
wwsch warz@aol. com

ABSTRACT

A revision of the dinematichthyine fishes (Ophidiiformes: Bythitidae: Brosmophycinae) of the Indo-West Pacific based
on more than 6500 specimens is published in several parts. Part IV is the last part and includes 4719 identified specimens
in the genera Alionematichthys (new genus with three described and eight new species), Dinematichthys Bleeker, 1855
(with one described and one new species) and Porocephalichthys (new genus with one described species). A neotype
of Dinematichthys iluocoeteoides Bleeker, 1855 is here designated. The genera reviewed here have in common a high
anterior nostril and are considered to be related to each other. When previously described, they were all included in
the genus Dinematichthys. The separating characters of the species are the pseudoclasper morphology, morphometric
characters, vertebrae and fin ray counts, otolith morphology, head squamation, presence or absence of the upper
preopercular pore and development of cirri on the snout.

Keywords: Viviparous brotulas, Indo-west Pacific, Dinematichthys, new species, new genera.

CONTENTS

INTRODUCTION.88

MATERIAL AND METHODS.88

COMPARATIVE MATERIAL.89

SYSTEM ATICS.89

Tribe Dinematichthyini Cohen and Nielsen, 1978 .90

Key to the genera and the species reviewed herein.89

Alionematichthys gen. nov.90

Alionematichthys ceylonensis sp. nov.92

Alionematichthys crassiceps sp. nov.95

Alionematichthys minyomma (Sedor and Cohen, 1987).99

Alionematichthys phuketensis sp. nov.100

Alionematichthys piger (Alcock, 1890).102

Alionematichthysplicatosurculus sp. nov.109

Alionematichthys riukiuensis (Aoyagi, 1954).113

Alionematichthys samoaensis sp. nov. 117

Alionematichthys shinoharai sp. nov.120

Alionematichthys suluensis sp. nov.122

Alionematichthys winterbottomi sp. nov.125

Alionematichthys sp. 1 .126

Alionematichthys sp. 2.127

Dinematichthys Bleeker, 1855.128

Dinematichthys iluocoeteoides Bleeker, 1855.129

Dinematichthys trilobatus sp. nov.136

Porocephalichthys gen. nov.138

Porocephalichthys dasyrhynchus (Cohen and Hutchins, 1982).138

DISCUSSION.141

REGIONAL CHECK LIST.142

KEY TO THE GENERA OF THE DINEMATICHTHYINI .144

ACKNOWLEDGMENTS.145

REFERENCES.145

W. Schwarzhans and P. R. Moller

INTRODUCTION

The global review of the dinematichthyine fishes, a tribe
within the subfamily Brosmophycinae of the viviparous
family Bythitidae, is concluded with this sixth part. The first
two parts dealt with the American Dinematichthyini (Moller
et al. 2004a, 2005) and were followed by four publications
revising the Dinematichthyini from the Indo-west Pacific
(Schwarzhans et al. 2005; Moller and Schwarzhans 2006;
Schwarzhans and Moller 2007; this paper). The latest
comprehensive account of ophidiiform fishes (Nielsen et
al. 1999) contained 12 genera now regarded as valid in the
Dinematichthyini, and 25 valid species. With the completion
of the world-wide review of the Dinematichthyini, the
current status of recognised taxa within the tribe has risen
to 26 genera with 110 species.

The fishes described in this paper were previously
referred to the genus Dinematichthys. Following Cohen
and Nielsen (1978) and redefined by Cohen and Hutchins
(1982) and Sedor and Cohen (1987), the position of the
anterior nostril high above the upper lip, and about midway
between the lip and the posterior nostril, was regarded as
the main diagnostic character for the genus, distinguishing
it from all other than known dinematichthyine fishes. In this
review we conclude that the formal genus Dinematichthys
“sensu Iato” in fact contains three well defined groups for
which generic rank applies: two more closely related ones -
Dinematichthys with two species and A lionematichthys gen.
nov. with 11 species - and one more distant monospecific
genus containing Pomcephalichthys gen. nov. dasyrhynchus
(Cohen and Hutchins). The combining character of the high
anterior nostril, reformulated to “positioned 1/3 or less of
the distance from upper lip to anterior margin of eye” in
Schwarzhans and Moller (2007), however, is not unique
to these three genera and this distinguishing character has
therefore become weakened. Similarly, high positioned
anterior nostrils between 1/3 and 1/3.5 the distance from
upper lip to anterior margin of eye have been observed in
a few species of the genus Diancistnis Ogilby (for instance
D. alleni, D. atollonm and D. katrineae), in Eusurculus
pistillum Schwarzhans and Moller and in Lapitaiclithys
frickei Schwarzhans and Moller. Fishes of the three
genera Dinematichthys, Alionematichthys gen. nov. and
Pomcephalichthys gen. nov. can be easily distinguished
from Diancistnis by the otolith sulcus showing distinctly
separated colliculi (vs fused, undivided), from Lapitaiclithys
in the presence of two pairs of pseudoclaspers (vs single
pair of outer pseudoclaspers), and from Eusurculus
pistillum in the flap-like inner pseudoclasper (vs stalked
with sucker-disk tip) (see also key below). The diagnostic,
distinguishing characters in the species of the three genera
covered here are similar to those of the previous parts of
our review. Again, pseudoclasper morphology represents
the most useful character. However, pseudoclaspers in
the following three genera are smaller than in most other
Dinematichthyini, with few exceptions, and arc also less

diverse in most instances. Other useful characters for
species differentiation are found to be head squamation,
vertebrae and fin ray counts, morphometric measurements,
otolith morphology, presence or absence of the upper
preopercular pore and, new in this group, development of
cirri on the snout. Of these, meristic counts show a lower
degree of variation than in most other Dinematichthyini, and
although individual species show statistically significant
differences (average values), they are rarely separated in
absolute values to be of use in diagnostic keys.

Eight species have been described previously, as placed
in the genus Dinematichthys, of which five are here regarded
as valid, and represent the three genera dealt with in this
part of the review. Three previously described species
are placed in synonymy. Dinematichthys Bleeker is the
earliest named genus of its tribe and hence, its definition is
essential to achieving nomenclatural stability. The genus
was established by monotypy (D. iluocoeteoides) based
on a single type specimen, which according to extensive
research by Cohen and Nielsen (1978) must be regarded
as no longer extant. For the purpose of redefinition of the
genus and species, another original Bleeker specimen
mentioned by Gunther (1862) (BMNH 1868.2.28.65),
probably the oldest extant, is here selected as neotype.
Although shrivelled, it represents a male specimen with
all the important diagnostic characters, and is in good
accordance with Blocker’s original description except for
a slight variation in eye size, which is within the range of
variability observed in the species.

MATERIAL AND METHODS

The examination of ca. 6500 specimens of Indo-west
Pacific Dinematichthyini yielded 4719 specimens which
were identified in the genera treated herein. Also included
are additional specimens identified in the collections of AMS
and USNM but not borrowed for detailed investigations.
These are listed as additional specimens and are not referred
to as type specimens for any of the new species.

The material described herein belongs to the following
institutions: AMS (Australian Museum, Sydney), ANSP
(Academy of Natural Sciences, Philadelphia), BMNH
(Natural History Museum, London), BPBM (Bishop
Museum, Honolulu), CAS (California Academy of
Sciences, San Francisco), KAUM (The Kagoshima
University Museum), KSHS (Kochi Prefectural Kochi
Nishi Senior High School, Kamobe, Kochi, Japan), MNHN
(Museum National d’Histoire Naturellc, Paris), NSMT
(National Science Museum, Tokyo), NTM (Museum and
Art Gallery of the Northern Territory (formerly Northern
Territory Museum), Darwin), ROM (Royal Ontario
Museum, Toronto), SAIAB (South African Institute for
Aquatic Biodiversity, formerly RUSI (JLB Smith Institute
of Ichthyology), Grahamstown), SMF (Senckenberg
Forschungsinstutut und Museum, Frankfurt/Main), SMNS

Dinematichthyine fishes of the Indo-west Pacific, Part IV

(Staatliches Museum fur Naturkunde, Stuttgart), TAU
(Tel Aviv University, Tel Aviv, Israel); USNM (National
Museum of Natural History, Smithsonian Institution,
Washington D.C.), WAM (Western Australian Museum,
Perth), YCM (Yokosuka City Museum), and ZMUC
(Zoological Museum, University of Copenhagen).

For methodology used in analysing dinematichthyine
fishes, reference is made to Moller et al. (2004a) and
Schwarzhans et al. (2005). Abbreviations used in meristic
counts are: D/V = anterior dorsal fin ray above vertebra
number; D/A = anterior anal fin ray below dorsal fin
ray number; V/A = anterior anal fin ray below vertebrae
number; D-A = number of dorsal fin rays minus number
of anal fin rays; V in D = number of dorsal fin rays per ray¬
bearing vertebra. The abbreviations ‘i.p.’ and ‘o.p.’ used in
the figures relating to male copulatory organ structures are
explained in Schwarzhans et al. (2005).

The ecology of most of the species is poorly known.
From available station data we have gathered some
information about habitat and depth range, but we have very
little data about behaviour, live colouration and feeding. A
number of females were examined for reproductive data,
e.g. number and size of embryos.

The distribution maps were created using Microsoft
Encarta 2001 digital world atlas.

COMPARATIVE MATERIAL

Indo-west Pacific Dincmatichthyini: see Schwarzhans
et al. (2005), Moller and Schwarzhans (2006) and
Schwarzhans and Moller (2007).

American Dinematichthyini: sec Moller et al. (2004a)
and Moller et al. (2005).

Brosmophycinae and Bythitinae: see Moller et al.
(2004b).

SYSTEMATICS

Family Bythitidac Gill, 1861
Subfamily Brosmophycinae Gill, 1862

Tribe Dinematichthyini Cohen and Nielsen, 1978

Diagnosis. Male copulatory organ with penis and 1-2
(rarely 3) pairs of pseudoclaspers in cavity of ventral body
wall covered by fleshy hood. First anal fin pterygiophore
slightly to strongly elongate. Head pore system generally not
reduced, 6 mandibular, 2-4 preopercular, 5-7 infraorbital
and 3-4 supraorbital pores, including supraorbital pore
above opercular spine. Posteriormost supraorbital head-
pore tubular.

Table 1 summarises meristic characters used in
distinguishing the species of the genera described here.

Key to the genera and the species reviewed herein

The following key to the genera Dinematichthys,
Alionematichthys gen. nov. and Porocephalichthys gen. nov.
is based on the key to the genera of the Dinematichthyini,

and on the species reviewed by Schwarzhans and Moller
(2007). The first dichotomy in that key refers to the anterior
nostril position and immediately separates the three genera
dealt with herein from all other dinematichthyine genera.
Since, however, certain other dinematichthyine genera exist
with almost similarly high anterior nostrils we begin the
following key with a separation from those few genera.

la. Anterior nostril positioned high (equal or less than
1/3 the distance from upper lip to anterior margin of

eye).2

1 b. Anterior nostril positioned low (1 /4 to 1/6 the distance

from upper lip to anterior margin of eye) .

all other Dinematichthyini (see Schwarzhans and
Moller (2007))

2a. Single pair of (outer) pseudoclaspers.

. Lapitaichthys

2b. Pair of inner and outer pseudoclaspers.3

3a. Sulcus of otolith with undivided colliculi.

. Diancistrus

[note: only a few species of the genus exhibit an
elevated anterior nostril]

3b. Sulcus of otolith with separated colliculi (not known
for Alionematichthys samoensis) .4

4a. Inner pseudoclasper stalked with sucker-disk shaped

tip. Eusurculus pistillum

[note: the only species in Eusurculus with an elevated
anterior nostril]

4b. Inner pseudoclasper variable, flap or stick-shaped or
folded, but not stalked and without sucker-disk shaped
tip.5

5a. Head with continuous squamation on cheeks, opercle

and occiput.6

[note: juveniles of Dinematichthys may have a
scaleless gap between cheeks and opercle]

5b. Head with scales on cheeks and in some species with
scale patch on opercle above opercular spine, rarely
also below opercular spine {Alionematichthys) .8

6a. Upper preopercular pore present, a pair of additional
mandibular pores, three additional pairs of pores on
occiput, one behind eye, two above preopercular pore;
sulcus of otolith with nearly equally long ostium and
cauda; precaudal vertebrae 14, total vertebrae 47^49,
dorsal fin rays 96-104, D/A 37-43; canine teeth

absent. Porocephalichthys dasyrhynchus

6b. Upper preopercular pore absent (see Schwarzhans
et al. 2005, fig. 1 A), no additional mandibular pores,
no pores on occiput; sulcus of otolith with ostium at
least two times as long as cauda; precaudal vertebrae
11-12, total vertebrae 41^45, dorsal fin rays 75-92,
D/A< 30; canine teeth present {Dinematichthys)...1

W. Schwarzhans and P. R. Moller

7a. Inner pseudoclasper with 2 lobes; head squamation

not extending beyond tip of maxilla.

. Dinematichthys iluocoeteoides

7b. Inner pseudoclasper with 3 lobes; head squamation
extending beyond tip of maxilla forward of 3rd

posterior mandibular pore.

. Dinematichthys trilohatus sp. nov.

8a. Upper preopercular pore present on both head sides;

cirri on snout present or absent.9

8b. Upper preopercular pore absent (rarely present on one
head side only); cirri on snout absent.15

9a. Cirri on snout absent, eye small (usually < 2% SL).
.10

9b. Cirri on snout present (except absent in Alionematichthys
suluensis sp. nov.), eye relatively large (usually > 2%
SL).11

10a. Scales above opercular spine 3-10; inner pseudoclasper
with anterior thorn as long as posterior lobe; otolith
length to otolith height 1.8-2.0, sulcus without
ventral indentation at junction of ostium and cauda
. Alionematichthys crassiceps sp. nov.

10b. Scales above opercular spine absent; inner
pseudoclasper with anterior thorn shorter than
posterior lobe; otolith length to otolith height 2.2-2.3,
sulcus with ventral indentation at junction of ostium
and cauda. Alionematichthys minyomma

11a. Scales above opercular spine 5-17; pscudoclaspers

not extruding beyond hood in resting position.12

[note: scales 2-6 in few tentatively assigned specimens
-Alionematichthys aff. riukiuensis ]

lib. Scales above opercular spine 0-2; pseudoclaspers
usually extruding beyond hood in resting position
.13

12a. Inner pseudoclasper thick, large, anterior thorn
entirely fused to anterior rim of outer pseudoclasper;
scale patch on opercle below opercular spine in large
specimens. Alionematichthys riukiuensis

12b. Inner pseudoclasper thin, small, bifurcate,
anterior thorn free from outer pseudoclasper;
no scale patch on opercle below opercular spine
. Alionematichthys winterbottomi sp. nov.

13a. Cirri on snout absent; anterior thorn of inner
pseudoclasper very long, twice as long as posterior
lobe. Alionematichthys suluensis sp. nov.

13b. Cirri on snout present; anterior thorn of inner
pseudoclasper about as long as posterior lobe.14

14a. Outer pseudoclasper with thick knob on inner face

distally; posterior nostril funnel-shaped.

. Alionematichthys samoaensis sp. nov.

14b. Outer pseudoclasper thin, without knob; posterior
nostril with anterior Hap, not funnel-shaped
. Alionematichthys piger

15a. Scales above opercular spine 2; sulcus of otolith
with extremely small cauda (ostium length to cauda
length 5.5-7, ostium height to cauda height 2.2-2.7)

. Alionematichthys shinoharai sp. nov.

15b. Scales above opercular spine absent; sulcus of otolith
with moderately small cauda (ostium length to cauda
length 3-4, ostium height to cauda height 1.5-2.0).
.16

16a. Inner pseudoclasper thick, with posterior lobe
folded over anterior thorn; outer pseudoclasper with

thickened tip...

. Alionematichthysplicatosurculus sp. nov.

16b. Innerpseudoclasperthin, unfolded; outer pseudoclasper
with thin or with knob on inner face.17

17a. Inner pseudoclasper with a symmetrically developed
anterior thorn and posterior lobe resembling in shape
a socket fitting to a small knob on the inner face of the
outer pseudoclasper; snout rounded; otolith length to

height 2.3-24, with shallow dorsal rim.

. Alionematichthys ceylonensis sp. nov.

17b. Inner pseudoclasper a simple flap without significant
indentation; snout pointed; otolith length to height

2.1-2.3, with distinct postdorsal projection.

. Alionematichthys phuketensis sp. nov.

Alionematichthys gen. nov.

(Tables 1-12)

Type species: Dinematichthys riukiuensis Aoyagi, 1954
(type locality: Ishigaki Island, Ryukyu Islands, Japan).
Gender masculine.

Dinematichthys (non Bleeker, 1855) in part. — Alcock
1890:432; Aoyagi 1954: 237; Cohen and Nielsen 1978: 57;
Sedorand Cohen 1987: 6; Machida 1994: 451; Nielsen et
al. 1999: 129; Moller el al. 2004a: 148.

Diagnosis. A genus of Dinematichthyini with the
following combination of characters: anterior nostril
placed high on snout at about one-third or less the distance
from lip to anterior rim of eye; head with a scale patch
on cheeks, in some species also above opercular spine,
rarely below opercular spine; male copulatory organ
with two pairs of mostly small pseudoclaspers, the outer
up to twice as large as inner; inner pseudoclasper with
anteriorly oriented pointed lobe containing supporter and
fleshy appendix posteriorly; large size, up to 150 mm SL;
precaudal vertebrae 10-12, total vertebrae 39-45, dorsal
fin rays 71-92, anal fin rays 55-72, V in D 2.0-24; upper
preopercular pore present or, less commonly, absent; sulcus
of otolith with ostium at least twice as long as cauda; maxilla
expanded posteroventrally.

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Table 1. Selected meristic characters of the species of Alionematichthys, Dinematichthys and Porocephalichthys: A, frequency distribution
of precaudal vertebrae, total vertebrae and anal fin ray counts; B, frequency distribution of dorsal fin ray and pectoral fin rays counts.

precaudal

vertebrae

total vertebrae

anal fin rays

Alionematichthys

revlonensis

Alionematichthys

crassiceps

Alionematichthys

minvomma

Alionematichthys

phuketensis

Alionematichthys

nicer

309

206

Alionematichthys

phcalosurculus

Alionematichthys

riukiuensis

Alionematichthys

samoaensis

Alionematichthys

shinolutrai

Alionematichthys

suluensis

Alionematichthys

winterhottomi

Alionematichthys

sp. 1

Alionematichthys

sp. 2

Dinematichthys

iluocoeteoides

147

Dinematichthys

trilobatus

Porocephalichthys

dasvrlivnchus

1V_

100

101

102

103

104

Alionematichthys

cevlonensis

Alionematichthys

crassiceps

Alionematichthys

minvomma

Alionematichthys

nliuketensis

Alionematichthys

nicer

Alionematichthys

nlicatosurculus

Alionematichthys

riukiuensis

Alionematichthys

samoaensis

Alionematichthys

shinolutrai

Alionematichthys

suluensis

Alionematichthys

winterhottomi

Alionematichthys

sp. 1

Alionematichthys

SEii

Dinematichthys

iluocoeteoides

Dinematichthys

trilobatus

Porocephalichthys

dasyrhynchus

W. Schwarzhans and P. R. Moller

Comparison. Alionematichthys is clearly related to
Dinematichthys, from which it mainly differs by lacking
a uniform and continuous head squamation, including the
cheeks, opercle and occiput (vs scale patches on the cheeks
and, in some species, on the opercle). The head squamation
of Dinematichthys (and Porocephalichthys gen. nov.) is
unique within the Dinematichthyini and it supports the
stability of generic diagnoses in the tribe. Dinematichthys
cannot easily be separated from Alionematichthys without
that character.

Species. Alionematichthys comprises II species; 10
from the Indo-west Pacific - A. ceylonensis sp. nov.,
A. crassiceps sp. nov., A. phuketensis sp. nov., A. piger
(Alcock, 1890) (formerly placed in Dinematichthys),
A. plicatosurcuhis sp. nov., A. riukiuensis (Aoyagi, 1954)
(formerly placed in Dinematichthys with Dinematichthys
megasoma Machida, 1994, as a junior synonym),
A. samoaensis sp. nov., A. sliinoharai sp. nov., A. suluensis
sp. nov., A. winterbottomi n. sp., and one species from the
tropical West Atlantic - A. minyomma (Sedor and Cohen,
1987) (formerly placed in Dinematichthys).

The species of the genus Alionematichthys can be
arranged in three species groups according to the presence
or absence of the upper preopercular pore, the presence
or absence of scales on the opercle, differences in the eye
size, and the presence or absence of cirri on the snout as
follows:

1) Alionematichthys plicatosurcuhis sp. nov. species
group: upper preopercular pore absent, no cirri on snout,
large eyes and no scales on the opercle, except for
A. shinoharai sp. nov. (two scales above opercular spine).
This group contains four species (A. ceylonensis sp. nov.,
A. phuketensis sp. nov., A. plicatosurcuhis sp. nov. and
A. shinoharai sp. nov.). However, strong differences in
the pseudoclasper and otolith morphology of some species
indicate that these species may not be closely related.

2) Alionematichthys riukiuensis species group: upper
preopercular pore present, numerous cirri on snout (except
cirri lacking in A. suluensis sp. nov.) and large eyes.
Presence or absence of scales on the opercle could be used to
separate two subgroups, one with five or more scales above
the opercular spine (A. riukiuensis and A. winterbottomi sp.
nov.), and the other usually without scales on the opercle
(rarely up to two scales above the opercular spine) (A. piger,
A. samoaensis sp. nov. and A. suluensis sp. nov.).

3) Alionematichthys minyomma species group: upper
preopercular pore present, no cirri on snout, small eyes and
scales on opercle present or absent (A. crassiceps sp. nov.
and A. minyomma). The small eyes and the big head readily
distinguish this small group, which combines a western
Pacific species and the only Atlantic species of the genus.

Remarks. Some of the more widespread species, namely
A. crassiceps sp. nov., A. piger and A. riukiuensis, show a
remarkable regionalisation usually based on variations of
a singie character.

Alionematichthys crassiceps sp. nov. and A. riukiuensis
contain few specimens, which can only be tentatively
assigned to the respective species. In the case of A. pigei-
the pseudoclaspers exhibit a wider than usual variation
which, however, does not seem to follow regional trends
in this particular case. The distinction of these (regional)
variations is not sufficient at this stage, but it is possible that
future genetic investigations may result in distinguishing
more species than we currently recognise.

Etymology. Combined from alius (Latin = the other,
different) and nematichthys, the stem of the genus name;
Dinematichthys, to which Alionematichthys is most similar.
The gender is masculine.

Alionematichthys ceylonensis sp. nov.

(Figs 1-3; Tables 1,2)

Material examined. (84 specimens, 19-72 mm SL),
HOLOTYPE-USNM 263707, male, 59 mm SL, Sri Lanka,
Weligama, 500 yards south of rest house, 05°57’25”N,
SO^fi’Q-’E^-bm.C. Koenig, 15 Feb. 1970. PARATYPES
- USNM 263731,4 males, 42-65 mm SL, 6 females, 47-63
mm SL, Sri Lanka, Weligama; USNM 366532,4 specimens,
37-72 mm SL, 08 o 33’24'’N, 81°14’42”E, Trincomalee, Sri
Lanka, 2-5 m, T. Iwamoto, 25 Sept. 2001; USNM 394975,
25 males, 37-69 mm SL and 40 females, 37-60 mm SL,
same data as holotype.

Additional material. USNM 366531, 7 specimens,
25-65 mm SL, Sri Lanka, Triconmalee; USNM 366533, 4
specimens, 40-59 mm SL, Sri Lanka, Battocalao; USNM
366534, 6 specimens, 34-60 mm SL. 07°56’N, 81°34'E,
Sri Lanka.

Diagnosis. Vertebrae 11-12+29-31=41-43, dorsal
fin rays 75-83 (average 79), anal fin rays 57-65 (average
60); eyes moderately large (2.0-2,8% SL); snout rounded,
without cirri; scales only on cheeks; no upper preopercular
pore; outer pseudoclasper broad-based, triangular, with
distal knob on inner side; inner pseudoclasper with
symmetrically developed anterior pointed lobe and
posterior lobe resembling in shape a socket fitting to small
knob on inner face of outer pseudoclasper; otolith length
to height 2.3-2.4, with shallow dorsal rim, otolith length
to sulcus length 1.5-1.6, ostium length to cauda length
3.3^L0.

Description (Figs 2-3).The principal meristic and
morphometric characters are shown in Tabic 2. Mature at
about 45 mm SL. Body slender, with rounded head profile.
No cirri on snout. Eye size 2.0-2.8 (2.6)% SL. Head with
scale patch on cheek containing 6-8 (8) vertical rows of
scales on upper part and 3 vertical rows on lower part.
Horizontal diameter of scales on body about 1.5% SL,
in 29 horizontal rows. Maxillary ending far behind eye,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded, with small knob. Anterior
nostril positioned high. 1/2.5 distance from upper lip to
anterior margin of eye. Posterior nostril small, about one-
fifth the size of eye. Opercular spine with free tip, thin and

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Fig. 1. Sample sites of O Alionematichthys ceylonensis sp. nov., © A. phuketensis sp. nov., © A. shinoharai sp. nov.. O A. plicatosurculus
sp. nov., © A. crassiceps sp. nov., and © A. aff. crassiceps. One symbol may represent several samples.

Fig. 2. Alionematichthys ceylonensis sp. nov., USNM 263707, holotype, male, 59 mm SL.

sharply pointed. Anterior gill arch with 14-18 (17) rakers,
3 elongate. A single short raker is placed between the two
lower elongate rakers in holotype and several paratypes.
Pseudobranchial filaments 2.

Head sensory pores (Fig. 3A, B). Supraorbital pores

2 to 3: first pore in front of second anterior infraorbital
pore, second pore above and behind eye usually absent,
third pore tubular, at upper termination of gill opening
above opercular spine. Infraorbital pores 6 (3 anterior and

3 posterior): first anterior pore behind anterior nostril,
second and third anterior pores covered by dermal flap of
upper lip, three posterior pores on rear part of upper lip,
about 1/3 the size of three anterior pores. Mandibular pores
6 (3 anterior and 3 posterior): first anterior pore large and
tubular, second anterior pore positioned in lateral skin fold,

small, third anterior pore at anterior termination of jugular
isthmus, three posterior pores on rear part of lower jaw.
Preopercular pores: 3 lower, first and second with joined
opening, covered by dermal flap in lateral view; third non¬
tubular; no upper preopercular pore. [This description of
the position of head sensory pores is used as reference for
all subsequent descriptions.]

Dentition (of holotype). Premaxilla with 6 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 1/4 diameter of pupil.
Vomer horseshoe-shaped, with 3 outer rows of small teeth
and I inner row of large teeth up to 1/5 diameter of pupil.
Palatine with 3 outer rows of small teeth and 1 inner row
of long teeth up to 1/4 diameter of pupil. Dentary with 5

W. Schwarzhans and P. R. Moller

Fig. 3. Alionematichthys ceylonensis sp. nov. A, lateral view of head, holotype; B, ventral view of head, holotype; C, ventral view of male
copulatory organ, holotype; D, view of left pscudoclaspers from inside, holotype; E, inclined lateral view of male copulatory organ, holotype;
F, view of left pscudoclaspers from inside, USNM 394975, 68 mm SL; G, view of left pseudoclaspers from inside, USNM 394975,48 mm
SL; H, median view of right otolith, USNM 394975, male, 58 mm SL.

outer rows of granular teeth and I inner row of larger teeth
anteriorly, up to about 1/3 diameter of pupil.

Otolith (Fig. 3H). Elongate in shape, length to height
2.3-2.4 (40-70 mm SL) and thin (otolith height to otolith
thickness about 2.5). Anterior and posterior tips moderately
pointed, the latter expanded and irregularly ornamented.
Dorsal rim shallow; ventral rim likewise shallow, deepest
anterior to middle. Inner face moderately convex, outer
face slightly concave, both smooth. Otolith length to sulcus
length 1.5-1.6. Sulcus slightly supramedialy positioned,
not inclined, with separated colliculi and marked notch at
ventral rim of sulcus at joint of ostium with cauda. Ostium
length to cauda length 3.3-4.0. Ventral furrow distinct, very
close to ventral rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined and
5-8 depressed. Parapophyses present from vertebrae 6 to
12(11). Pieural ribs on vertebrae 2 to 11 (10). First anal fin

pterygiophore not or only slightly longer than subsequent
pterygiophore.

Male copulatory organ (Fig. 3C-G). Two pairs of
moderately large pseudoclaspers, outer pair about 50%
larger than inner; outer pseudoclasper with broad base,
triangular with pointed tip and distinct knob at inner face
near tip; inner pseudoclasper with symmetrically developed
anterior thorn and posterior lobe resembling in shape a
socket fitting to small knob on the inner face of the outer
pseudoclasper. Isthmus narrow; penis curved, about as long
as outer pseudoclaspers.

Colour. Live colour unknown. Preserved colour
uniformly dark brown.

Comparison. Alionematichthys ceylonensis belongs
to the species group without an upper preopercular pore
together with A. phuketensis sp. nov., A. plicatosurculus
sp. nov. and A. shinoharai sp. nov., from which it differs
in the peculiar shape of the inner pseudoclasper and the

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Table 2. Meristic and morphometric characters of Alionematichthys
ceylonensis sp. nov.

Holotype +
Holotype 83 paratypes
USNM-
263707

Mean (range)

Standard length in mm

53.9(19-72)

Meristic characters

Dorsal fin rays

78.9 (75-83)

Caudal fin rays

15.9(15-17)

Anal fin rays

60.5 (57-65)

Pectoral fin rays

20.9(19-22)

Precaudal vertebrae

12 . 0 ( 11 - 12 )

Caudal vertebrae

30.0 (29-31)

Total vertebrae

42.0 (41-43)

Rakers on anterior gill arch

16.6(14-18)

Pseudobranchial filaments

D/V

5.8 (5-6)

D/A

22.6 (20-26)

V/A

13.3(13-15)

Morphometric characters in % of SL

Head length

27.8

26.8 (25.8-27.8)

Head width

15.3

14.1 (12.1-16.1)

Head height

16.0

16.4(14.8-17.9)

Snout length

5.5

6.0 (5.4-6. 8 )

Upper jaw length

14.8

11.8-15.1 (14.0)

Diameter of pigmented eye

2.6

2.5 (2.0-2. 8 )

Diameter of pupil

1.1

1.5(1.1-1.7)

Interorbital width

7.8

7.3 (6.3-8.0)

Posterior maxilla height

4.7

4.4 (3,44.9)

Postorbital length

19.9

19.4(18.5-20.7)

Preanal length

47.9

47.5(45.8-51.7)

Predorsal length

32.4

31.0(30.0-32.5)

Body depth at origin of anal

18.7

18.3(14.5-20.1)

fin

Pectoral fin length

12.6

12.6(11.7-13.2)

Pectoral fin base height

6.5

6.2 (5.2-7.2)

Ventral fin length

23.3

22.7(19.2-24.5)

Base ventral fin - anal fin origin

28.3

28.3 (26.7-32.8)

knob at the inner face of the outer pseudoclasper and the
slender otoliths (otolith length to otolith height 2.3-2.4 vs
2.1-2.3). From A. phuketensis sp. nov., it differs further
in the rounded snout profile (vs pointed) and the lack
of a distinct postdorsal projection of the otolith. From
/(. shinoharai sp. nov.. it further differs in the lack of scales
above the opercular spine (vs 2), the long sulcus (otolith
length to sulcus length 1.5-1.6 vs 1.9-2.0) and the notch at
the ventral sulcus margin at the joint of ostium and cauda
(vs no notch).

Distribution. Alionematichthys ceylonensis occurs only
along the shores of the island of Sri Lanka (Fig. 1).

Biology. A 53 mm SL gravid female (USNM 394975)
contains 80 embryos, length 4.8 mm TL, and 60 orange
eggs, diameter 0.7 mm.

Etymology. Named after the type locality, Sri Lanka,
previously known as Ceylon. It is an adjective.

Alionematichthys crassiceps sp. nov.

(Figs 1,4-6; Tables 1,3)

Material examined. 53 specimens, 34-63 mm SL.
HOLOTYPE - BPBM 9310, male, 63 mm SL, Truk,
Caroline Islands, Southfield beach, 08°0’N, 147°0’E,
inshore, 0-4 m, rock and Halimeda algae, J. E. Randall, 10
July 1969. PARATYPES - BPBM 9217, 1 male, 40 mm
SL, 1 female, 42 mm SL, Palau Islands, NE Arakabesan
Island, 0.5-1 m, corals, J. E. Randall and E. S. Helfman,
5 June 1968; BPBM 40932, 3 females, 46, 46 and 47 mm
SL, 3 males, 34, 36,43 mm SL, same data as for holotype;
AMS I. 20547-042, 1 male, 40 mm SL, 2 females, 45 and
48 mm SL, Panasesa reef, Louisiadc Archipelago, Papua
New Guinea, 5-8 m, B. Goldman, 24 March 1969; CAS
227282, 1 male, 49 mm SL, 1°4’28”N, 154°42 , 28”E,
0-15 m, lagoon coral heads on inner margin of reef flat at
Tewawaelal, extreme W end of atoll; low tide; water clear,
sand & coral. Kapingamarangi Atoll, Pohnpei, Caroline
Islands, Micronesia, Atta, Kindaro and H. H. Rofen, 14
July 1954; CAS 222543,2 males, 50 and 52 mm SL, Vanua
Levu, Fiji; CAS 222563, 2 males, 40 and 47 mm SL, 1
female, 37 mm SL, Suva, Fiji; CAS 227285, 1 female,
52 mm SL, Vanua Levu, Fiji; CAS 227303, 1 female, 30
mm SL, 1 male, 40 mm SL, Suva, Fiji; USNM 223247,
1 male, 55 mm SL, 07°00’30”N, 158°11 , 55”E, Caroline
Islands, Pohnpei, off east side of Jokaj Passage, 0-4 m, V. G.
Springer and party, 5 Sept. 1980; USNM 394976,3 females,
42, 43 and 56 mm SL, 4 males 40-57 mm SL, 2 juveniles
12 and 22 mm SL, 18°44’S, 174°06’W, Vava’u Islands,
Tonga, 0-11 m, J.T. Williams et ai, 17 Nov. 1993; USNM
352743, 7 males, 48-60 mm SL, 5 females, 47-67 mm
SL, 18°44’25'’S, 169° 12’41”E, Vanuatu, Erromango, Port
Narevin, 0-5 m, J. T. Williams et al., 28 May 1996; USNM
374196, 1 male, 57 mm SL, 01°3ri2”S, 145°01’30”E,
Papua New Guinea, Hermit Islands, Jalun Island, V. G.
Springer and party, I Nov. 1978.

Tentatively assigned specimens. USNM 366687, 3
males, 47,50 and 53 mm SL, Bay SE of K’en Ting, Taiwan,
0-3 m, V. G. Springer and party, 22 April 1968.

Additional material. USNM 334123, 1 male, 49 mm
SL, Tonga; USNM 374210, 1 male, 45 mm SL, 4 females,
34-50 mm SL, Tongatapu, Tonga.

Diagnosis. Vertebrae 10-11+29-31=40-42, dorsal fin
rays 72-81, anal fin rays 57-64; eyes small (1.3-2.5% SL);
snout blunt, without cirri; scales on cheeks and patch above
opercular spine with 3-10 scales; upper preopercular pore
present; outer pseudoclasper broad-based, simple flap like;
inner pseudoclasper with free, forward-pointing anterior
lobe and broad posterior lobe; otolith length to height
1.9-2.0, with rounded dorsal rim, otolith length to sulcus
length 1.7-1.8, cauda very short, ostium length to cauda
length 4.5-5.0.

W. Schwarzhans and P. R. Moller

Fig. 4. Alionematichthys crassiceps sp. nov., BPBM 9310, holotype, male, 63 mm SL.

Description (Figs 4-6).The principal meristic and
morphometric characters are shown in Table 3. Mature at
about 45 mm SL. Body stout, with rounded, robust head
profile (interorbital width 6.6-8.9 (7.4)% SL). No cirri on
snout. Eye size 1.3-2.5 (1.3)% SL. Head with scale patch
on cheek containing 6-10(10) vertical rows of scales on
upper part and 3-5 vertical rows on lower part. Horizontal
diameter of scales on body of holotype 2.1% SL, in about
26 horizontal rows. Maxillary ending far behind eye, dorsal
margin of maxillary covered by upper lip dermal lobe,
posterior end strongly expanded. Anterior nostril positioned
high, one-third distance from upper lip to anterior margin of
eye. Posterior nostril small, about one-sixth the size of eye
or less. Opercular spine with free tip, pointed. Anterior gill
arch with 16-19 (16) rakers, 3 elongate. Pseudobranchial
filaments 1-2 (1).

Head sensory pores (Figs 5A-E, 6A,B). Supraorbital
pores 2 to 3. Infraorbital pores 6 (3 anterior and 3 posterior),
posterior pores about one-third the size of anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior): first anterior
pore large, tubular, with cirrus. Preopercular pores: 3 lower,
first and second with joined opening, covered by dermal flap
in lateral view; third nontubular; small upper preopercular
pore. [See description of Alionematichthys ceylonensis for
position of pores.]

Dentition (of holotype). Premaxilla with 6 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriomiost teeth in inner row up to 3/4 diameter of pupil.
Vomer horseshoe-shaped, with 4 outer rows of small teeth
and 1 inner row of large teeth up to 1/2 diameter of pupil.
Palatine with 4 rows of equally long teeth up to I /2 diameter
of pupil. Dentary with 2 outer rows of granular teeth and
2 inner rows of larger teeth anteriorly, up to about size of
diameter of pupil.

Otolith (Figs 5M-P, 6E). Compact in shape, length to
height 1.9-2.0, moderately thin (otolith height to otolith
thickness about 2.5). Anterior tips moderately pointed,
posterior tip much expanded postero-dorsally. Dorsal rim
evenly curved; ventral rim deep, deepest at its middle or
slightly anteriorly. Inner face moderately convex, outer face
flat, both smooth. Otolith length to sulcus length 1.7-1.8.
Sulcus mcdianly positioned, not inclined, with separated
colliculi but no notch at ventral rim of sulcus at joint of

ostium with cauda. Cauda very small, ostium length to
cauda length 4.5-5.0. Ventral furrow distinct, very close
to ventral rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined and
5-8 depressed. Parapophyscs present from vertebrae 6 to
11 (10). Pleural ribs on vertebrae 2 to 10 (9). First anal fin
pterygiophore not or only slightly longer than subsequent
pterygiophore.

Male copulatoiy organ (Figs 5F-L, 6C, D). Two pairs
of moderately large pscudoclaspers, outer pair about 50%
larger than inner; outer pseudoclasper with widened base
and rounded tip, fiat, flap-like; inner pseudoclasper with
free, forward-pointing anterior thorn, connected at base
to anterior rim of outer pseudoclasper and broad posterior
lobe with deep inclined furrow on its inner margin. Isthmus
moderately wide; penis curved, somewhat longer than outer
pseudoclaspers.

Colour. Live colour unknown. Preserved colour
uniformly light brown.

Remarks. Alionematichthys crassiceps shows a high
degree of regional differentiation.

Specimens from Vanuatu (Fig. 5C) and Fiji have a
generally higher number of scales (7-10) above the opercular
spine than those from Tonga (Fig. 5D) or Micronesia (Fig.
5A, B) (5-7) on either side of the distribution range. Those
from Tonga show a peculiar pattern in the way that 3-4
larger scales form a straight row with 2-3 small scales
above (Fig. 5D). The ones from western Micronesia (Truk
and Palau) (Fig. 5A-B) have the highest number of vertical
scale rows on the cheeks (mostly 8 vs 6-7). Those from
Pohnpei and Kapingamarangi Atoll (Fig. 5E) appear to
have slightly larger eyes than those from other locations.
All these differences, however, are very subtle and are not
supported by other characters so as to warrant a separation
into different species.

Three specimens from Taiwan (Fig. 6A-E) are
tentatively placed in the species (A. afif. crassiceps ) due to
some folded, cirri-like dermal flaps on the chin, a generally
lower number of scales above the opercular spine (3-6), a
rather short anterior thorn of the inner pseudoclasper and its
remote location from the other specimens of the species.

Comparison. Alionematichthys crassiceps is readily
recognised by its big head and the small eyes, both characters

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Fig. 5 (part). Alionematichthys crassiceps sp. nov.. A, lateral view of head, holotype; B, ventral view of head, holotype; C, lateral view
of head, holotype; D, lateral view of head, USNM 394976, male, 49 mm SL; E, lateral view of head, USNM 223247, male, 55 mm SL; F,
inclined lateral view of male copulatory organ, holotype; G, ventral view of male copulatory organ, USNM 374196, 58 mm SL; H, view
of left pseudoclaspers from inside, holotype; I, view of left pseudoclaspers from inside, USNM 394976, male, 49 mm SL; ,J, view of left
pseudoelaspers from inside, USNM 394976, 57 mm SL; K, view of left pseudoclaspers from inside, CAS 222543,50 mm SL; L, view of left
pseudoclaspers from inside, USNM 223247, 55 mm SL.

W. Schwarzhans and P. R. Moller

Fig. 5 (continued). M, median view of right otolith, CAS 222543, male, 50 mm SL; N, median view of right otolith, USNM 352743, female, 67
mm; O. median view of right otolith, USNM 374196, male, 58 mm SL; P, median view of right otolith, USNM 394976, female, 56 mm SL.

that connects it with the West Atlantic A. minyomina, from
which it is distinguished by the presence of scales above
the opercular spine (vs absent) and the compact otolith
with an otolith length to otolith height ratio of 1.9-2.0 (vs
2.2-2.3). The only co-occurring Dinematichthyini with big

heads and small eyes are those of the genus Diancistms
Ogilby of the Diancistms erythraeus species group which,
however, shows a completely different pseudoelasper
pattern (see Schwarzhans et ctl. 2005), no scales above the
opercular spine and an otolith with an undivided sulcus

Fig. 6. Alionematichthys aff. crassiceps, USNM 366687, A, lateral view of head; B, ventral view of head; C, inclined lateral view of male
copulatory organ; D, view of left pseudoclaspers from inside; E, median view of right otolith.

Dinematichthyine fishes of the Indo-west Pacific, Part IV

and fused colliculi. Front other co-occurring species of
Alionematichthys, it is distinguished by the following
additional characters: from A. piger, by the small outer
pseudoclasper (vs long), the lack of cirri on the snout (vs
present) and the presence of scales above the opercular
spine (vs absent); from A. plicatosurculus sp. nov., by the
different pseudclasper pattern, the presence of an upper
preopercular pore (vs absent), the presence of scales above
the opercular spine (vs absent); from A. riukiuensis, through
the anteriorly free inner pseudoclasper (vs connected), the
compressed otolith (vs elongate) and the lack of cirri on the
snout (vs present); and from A. winterbottomi sp. nov., in
the shape of the inner pseudoclasper and the lack of cirri
on the snout (vs present).

Distribution. Alionematichthys crassiceps occurs along
oceanic islands of the West Pacific from Micronesia (Palau,
Chuuk, Pohnpei, Kapingamarangi Atoll), Hermit Island and

Table 3. Meristic and morphometric characters of Alionematichthys
crassiceps sp. nov.

Holotype

BPBM

9310

Holotype +

42 paratypes

Mean (range)

Standard length in mm

46.6(12-67)

Meristic characters

Dorsal fin rays

76.0(72-81)

Caudal fin rays

Anal fin rays

60.7 (57-64)

Pectoral fin rays

21.7(20-23)

Precaudal vertebrae

11.0(10-11)

Caudal vertebrae

30.2(29-31)

Total vertebrae

41.2(40-42)

Rakers on anterior gill arch

17.4(16-19)

Pseudobranchial filaments

1.9 (1-2)

D/V

5.9 (5-7)

D/A

20.6(18-23)

V/A

12.7(12-13)

Morphometric characters in % of SL

Head length

28.1

30.0(28.1-34.3)

Head width

17.6

15.6(13.7-18.1)

Head height

19.9

18.6(17.3-20.8)

Snout length

7.2

6.7 (5.7-8.0)

Upper jaw length

14.7

14.5(13.6-16.9)

Diameter of pigmented eye

1.3

1.9(1.3-2.5)

Diameter of pupil

1.0

1.2 (0.9-1.8)

Interorbital width

7.4

7.8 (6.6-8.9)

Posterior maxilla height

4.7

4.9 (4.6-5.7)

Postorbital length

21.0

22.3 (21.0-26.3)

Preanal length

49.1

49.1 (44.0-57.9)

Predorsal length

33.1

33.1 (31.6-36.6)

Body depth at origin of anal fin

20.6

19.4(17.9-23.1)

Pectoral fin length

15.5

15.5(12.6-18.8)

Pectoral fin base height

6.6

6.7 (6.0-8.2)

Ventral fin length

21.8

26.3(21.8-29.8)

Base ventral fin - anal fin origin

25.7

27.7(24.3-32.1)

the Kiriwina (Trobriand) Islands of New Guinea, Vanuatu,
Fiji to Tonga (Fig. 1). There are a few tentative records
from Taiwan.

Biology. A 56 mm SL female (USNM 394976)
contained 95 embryos, length 4.1 mm TL, and ca. 50 orange
eggs, diameter 0.6 mm.

Etymology. Named after the big head characteristic
for the species; crassus (Latin = thick) and adjective
ending derived from caput (Latin = head). It is a noun in
apposition.

Alionematichthys minyomma (Sedor and Cohen, 1987)

(Figs 7, 8; Tables 1,4)

Dinematichthys minyomma Sedor and Cohen, 1987:
6; Nielsen et al. 1999: 130; Moller et al. 2004: 149 (as
D. minyomma).

Species 2. — Sedor 1985.

Tabic 4. Meristic and morphometric characters of Alionematichthys
minyomma (Sedor and Cohen, 1987)

Holotype

USNM

280122

Holotype +

10 paratypes
and

28 non-types
Mean (range)

Standard length in mm

53.3 (26-78)

Meristic characters

Dorsal fin rays

74.7 (71-79)

Caudal fin rays

Anal fin rays

57.8 (55-61)

Pectoral fin rays

21.7(21-23)

Precaudal vertebrae

10.8(10-11)

Caudal vertebrae

29.1 (29-30)

Total vertebrae

39.8 (39-40)

Rakers on anterior gill arch

17.5(15-19)

Pseudobranchial filaments

D/V

5.5 (5-6)

D/A

22.4 (21-25)

V/A

13.1 (12-14)

Morphometric characters in % of SL

Head length

29.0

28.0 (26.6-29.8)

Head width

17.0

15.5(13.8-17.5)

Head height

23.5

20.3 (18.0-23.7)

Snout length

7.0(6.3-7.9)

Upper jaw length

15.5

14.8(13.8-15.9)

Diameter of pigmented eye

1.5

2.0(1.6-2.3)

Diameter of pupil

1.2 (0.9-1.6)

Interorbital width

8.5

8.1 (7.0-9.1)

Posterior maxilla height

6.0

5.0 (4.8-54)

Postorbital length

22.0

20.3(19.3-21.5)

Preanal length

49.5

49.9 (45.4-56.0)

Predorsal length

32.5

32.1 (30.7-34.0)

Body depth at origin of anal fin

20.9(18.9-23.4)

Pectoral fin length

13.5

14.8(13.3-16.8)

Pectoral fin base height

7.1 (6.6-8.1)

Ventral fin length

28.0

25.6 (20.7-28.5)

Base ventral fin - anal fin origin

32.0

29.1 (25.9-32.1)

W. Schvvarzhans and P. R. Moller

110*W 100*W 90"W 80*W 70*W 60'W 50'W

Fig. 7. Sample sites of O Alionematichthys minyomma (Sedor and
Cohen, 1987). One symbol may represent several samples.

Material examined. (40 specimens, 26-80 mm SL):
see Moller et al., 2004.

Diagnosis. Vertebrae 10-11+29-30=39-40, dorsal fin
rays 71-79. anal fin rays 55-61; eyes small (1.5-2.3% SL);
snout rounded, without cirri; scales only on cheeks; upper
preopercular pore present; outer pseudoclasper broad-
based, simple flap-shaped, about twice as large as inner
pseudoclasper; inner pseudoclasper very short, its margin
almost straight from short anterior pointed lobe to posterior
lobe; isthmus between pseudoclaspers very wide; otolith
length to height 2.2-2.3, with regular dorsal rim, otolith
length to sulcus length 1.6, ostium length to cauda length
about 3.5-4.0.

Remarks. For a detailed description and additional
illustrations see Moller et al. (2004) (as Dinematichthys
minyomma).

Comparison. Alionematichthys minyomma belongs to
the small species group of the genus characterised by their
small eyes and big head, further characterised by the lack
of cirri and the presence of an upper preopercular pore. It
is distinguished from the only other species of the group
(A. crassiceps ) by the lack of scales above the opercular
spine, differences in the shape of the inner pseudoclaspers
and the slender otoliths (otolih length to otolith height
2.2-23 vs 1.9-2.0).

Distribution. Alionematihthys minyomma represents
the only species of the genus in the western tropical Atlantic
(off Honduras, Atlantic Panama and Columbia, Haiti,
Puerto Rico and northern Antilles) (Fig. 7) i.e. outside of
the Indo-west Pacific.

Alionematichthysphuketensis sp. nov.

(Figs 1, 10, 11, Tables 1,5)

Material examined. (9 specimens, 25-41 mm SL).
HOLOTYPE - ZMUC P771659, male 40 mm SL, Rawai
beach, Phuket, Thailand, 07°47’0”N, 98° 19’0”E, J. Nielsen,
24 Nov. 1972. PARATYPES-ZMUC P771660-67, 1 male,
36 mm SL, 6 females, 28^H mm SL, 1 juvenile, 25 mm
SL, same data as holotype.

Diagnosis. Vertebrae 12+30-31=42^43, dorsal fin rays
81-89, anal fin rays 60-65; eyes moderately large (2.3-
2.9% SL); body slender, snout pointed, without cirri; scales
only on cheeks; no upper preopercular pore (occasionally
single pore on one side); outer pseudoclasper broad-based,
small; inner pseudoclasper small, anteriorly inclined flap;
otolith length to height 2.1-2.3. with distinct postdorsal

Fig. 8. Alionematichthys minyomma (Sedor and Cohen, 1987). A. lateral view of head, USNM 280123, male, 66 mm SL; B, ventral view of
head, USNM 280123, female, 61 mm SL; C, view of left pseudoclaspers from inside, USNM 280123, male, 71 mm SL; D, inclined lateral
view of male copulatory organ of same specimen; E, median view of right otolith, USNM 280123, male 71 mm SL.

100

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Table 5. Meristic and morphometric characters of Alionematichthys
phuketensis sp. nov.

Holotype

ZMUC

P771659

Holotype +

8 paratypes

Mean (range)

Standard length in mm

Meristic characters

33.0(25-41)

Dorsal fin rays

83.3 (81-89)

Caudal fin rays

15.2(14-16)

Anal fin rays

62.9 (60-65)

Pectoral fin rays

19.9(19-21)

Precaudal vertebrae

Caudal vertebrae

30.8 (30-31)

Total vertebrae

42.8 (42-43)

Rakers on anterior gill arch

16.0(15-18)

Pseudobranchial filaments

D/A

23.1 (21-25)

V/A 14

Morphometric characters in % of SL

13.3(13-14)

Head length

26.8

26.7(25.1-27.5)

Head width

12.5

11.7(11.0-12.5)

Head height

15.0

15.1 (14.0-16.1)

Snout length

6.2

6.2 (5. 6 - 6 .5)

Upper jaw length

13.1

12.9 (12.5-13.7)

Diameter of pigmented eye

2.9

2.6 (2.3-2.9)

Diameter of pupil

1.8

1.6 (1.4-1. 8 )

Interorbital width

7.3

7.3 ( 6 .9-7.9)

Posterior maxilla height

4.4

3.7 (3.3-4.4)

Postorbital length

18.4

18.6(17.9-19.4)

Preanal length

44.9

43.4 (41.0-45.6)

Predorsal length

31.0

30.7 (29.2-32.2)

Body depth at origin of anal fin

16.9

15.3(13.7-16.9)

Pectoral fin length

12.4

13.2(11.9-15.0)

Pectoral fin base height

5.2

5.8 (5.2-6.3)

Ventral fin length

19.8

23.5 (19.8-24.8)

Base ventral fin - anal fin origin

25.2

23.9 (22.4-25.2)

projection, otolith length to sulcus length 1.6-1.7, ostium
length to cauda length 3.0-3.3.

Description (Figs 10, llj.The principal meristic and
morphometric characters are shown in Table 5. Mature
at about 40 mm SL (largest male at 40 mm SL probably
subadult). Body slender, with pointed head profile. No cirri
on snout. Eye size 2.3-2.9 (2.9)% SL. Head with scale patch
on check containing 7 vertical rows of scales on upper part
and 4 vertical rows on lower part. Horizontal diameter
of scales on body about 1.5% SL, in about 30 horizontal
rows. Maxillary ending far behind eye, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior end
expanded. Anterior nostril positioned high, 1/3 distance
from upper lip to anterior margin of eye. Posterior nostril
small, about 1/8 the size of eye. Opercular spine with free
tip, thin and sharply pointed. Anterior gill arch with 15-18
(18) rakers, 3 elongate. In holotype and two paratypes,

lower two elongate rakers interrupted by one plate-like
raker. In rest of paratypes, three elongate rakers in single
row. Pseudobranchial filaments 2.

Head sensoiy pores (Fig. 11 A, B). Supraorbital pores
2. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about 1/3 the size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior): first anterior
pore large, tubular, without cirri. Preopercular pores: 3
lower, first and second with joined opening, covered by
dermal flap in lateral view; third non-tubular; no upper
preopercular pore (rarely single pore on one side). [See
description of Alionematichthys ceylonensis for position
of pores.]

Dentition (of holotype). Premaxilla with 5 outer rows
of granular teeth and I inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 1/4 diameter of pupil.
Vomer horseshoe-shaped, with I outer row of small teeth
and 1 inner row of large teeth up to 1/5 diameter of pupil.
Palatine with 3 outer rows of small teeth and 1 inner row
of large teeth up to 1/5 diameter of pupil. Dentary with 5
outer rows of granular teeth and 1 inner row of larger teeth
anteriorly, up to about 1/3 diameter of pupil.

Otolith (Fig. 11E-G). Elongate in shape, length to height
2.1-2.3 (36-41 mm SL) and moderately thin (otolith height
to otolith thickness about 1.6). Anterior tip moderately
pointed, posterior tip expanded and irregularly ornamented.
Dorsal rim shallow, with distinct postdorsal projection and
furrow thereafter. Inner face moderately convex, outer face
almost flat, both smooth. Otolith length to sulcus length
1.6-1.7. Sulcus slightly supramedially positioned, slightly
inclined, with separated colliculi and marked notch at
ventral rim of sulcus at joint of ostium with cauda. Ostium
length to cauda length 3.0-3.3. Ventral furrow very close
to ventral rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined
and 5-8 depressed. Parapophyses present from vertebrae
6 to 12. Pleural ribs on vertebrae 2 to 11. First anal fin
pterygiophore not or only slightly longer than subsequent
pterygiophore.

Male copulatory organ (Fig. 11C, D). Two pairs of
small pseudoclaspers, which may not be fully mature in
all specimens investigated. Outer pseudoclasper a simple
flap with broad base; inner pseudoclasper a forward
inclined small flap without significant indentation. Isthmus
moderately narrow; penis curved, longer than outer
pseudoclaspers.

Colour. Live colour unknown. Preserved colour
uniformly light brown.

Comparison. Alionematichthys phuketensis belongs
to the species group without an upper preopercular pore
together with A. ceylonensis, A. plicatosurculus sp. nov.
and A. shinoharai sp. nov. It differs from all of them in
the combination of a slender body shape with a pointed
snout and the peculiar postdorsal projection of the otolith
rim. From A. plicatosurculus sp. nov., it differs further
in the simple inner pseudoclasper morphology and from

101

W. Schwarzhans and P. R. Moller

90°E 100*E 110'E 120'E 130*E 140*E 150*E 160*E 170’E 180* 170’W 160*W 150*W 140*W 130°W 120*W

Fig. 9. Sample sites of © Alionematichthys suhiensis sp. nov. and O A. piger (Alcock. 1890). One symbol may represent several samples.

A. shinoharai sp. nov., in the lack of scales above the
opercular spine (vs two), the longer sulcus (otolith length
to sulcus length 1.6-1.7 vs 1.9-2.0) and the notch at the
ventral sulcus margin at the joint of ostium and cauda (vs
no notch). Alionematichthys phuketensis co-occurs with
A. piger, A. nukiuensis and Dinematichthys iluocoeteoides.
From the latter two it differs readily in the absence of scales
above the opercular spine (vs present or entire operclc with
scales in D. iluocoeteoides). From A. piger it di tiers in the
lack of cirri on the snout, the lack of an upper prcopcrcular
pore and the peculiar dorsal projection of the otolith rim.

Distribution. Alionematichthys phuketensis so far
is only known from the type locality, Ravai Beach near
Phuket, Thailand (Fig. I).

Etymology. Named after the type locality, the town of
Phuket, Thailand. It is an adjective.

Alionematichthys piger (Alcock, 1890)

(Figs 9, 12, 13, Tables 1,6)

Dinematichthys piger Alcock, 1890: 432 (Great Coco
Island, Andaman Islands); Alcock, 1905: plate 37, fig. 3;

IBrotulina piger (Alcock, 1890). — Nielsen el al.
1999: 126.

Material examined. 2288 specimens, 16-84 mm SL.
AMS 1.17094-070, 3 males, 5 females, 11 juveniles,
08°30’S, 151°05’E, Kiriwina Islands (Trobriand), Papua
New Guinea; AMS 1.17102-004, 2 specimens, 08°30’S
I51°05’E, Kiriwina Islands (Trobriand), Papua New
Guinea; AMS 1.17424-012, 1 female, 31 °32’S, 159°04’E,
Lord Howe Island; AMS 1.18052-040, 2 females, 28-30
mm SL, 01°44’N, 172°59’E, AbaiangAtoll, Gilbert Islands,
Kiribati; AMS 1.20629-001, 3 males, 10 females, 12°N,

165°E, Bikini Atoll, Marshall Islands; AMS 1.27134-040,
1 female, 29°27’2”S, 159°06’8”E, Lord Howe Island, 6-9
m; AMS 1.27138-047, 1 male, 46 mm SL. 2 females, 34-67
mm SL,29°27'8”S, 159°05'E, Middleton Reef, Lord Howe
Rise, Tasman Sea, 4-9 m; AMS 1.27148-029, 1 female,
29°57'S, 159°01’2”E, Elizabeth Reef, Tasman Sea, 0-10
m; AMS 1.27891-040, 1 specimen, 29°56’S, I59°01’E,
Elizabeth Reef, Tasman Sea, 0-5 m; AMS 1.28950-045, 2
specimens, 17°29’1”S, 149°5r3”W, Moorea, French
Polynesia, 8-12 m; AMS 1.33740-026,3 males, 49-59 mm
SL and 6 females, 10°09.58”S, 144°34.94”E, Ashmore
Reef, Coral Sea; AMS 1.34510-020, 42 specimens,
15°44.8’S, 144°38.9'W,Taiaro Atoll,Tuamotu Islands, 0-2
m; AMS 1.37315-032, 2 males, 65-75 mm SL, 4 females,
40-54 mm SL, 18°44'45”S, 169°12’68”E, Erromango,
Vanuatu, 0-6 m; AMS 1.39387-001,12 specimens, Pelagai
Island, Guam, 1-2 m; BPBM 7553, 1 male, 4 females,
Fanning Island, Line Islands; BPBM 8009, 2 males, 5
females, Marshall Islands; BPBM 8242, 1 male, 2 females,
Enewetak Atoll, Marshall Islands; BPBM 8651, 1 male, 5
females, Tahiti; BPBM 8658, I female, Tahiti; BPBM 9311,
1 female, Chuuk, Micronesia; BPBM 10240, 1 specimen,
Rangiroa, Tuamotu Islands; BPBM 1 0307, 2 specimens,
Rangiroa, Tuamotu Islands; BPBM 10658, 1 male and 9
females, Gilbert Island, Kiribati; BPBM 10830,1 specimen,
Aitutaki, Cook Islands; BPBM 11588, 1 female, Tahiti;
BPBM 12254, 1 male, 2 juveniles, Ducie, Pitcairn; BPBM
13574, 1 male and 1 female, Mangareva, Gambier Islands,
French Polynesia; BPBM 13915, 2 females, Rarotonga,
Cook Islands; BPBM 13929, I male. Rarotonga, Cook
Islands; BPBM 14161, 1 female. Palmyra Atoll, Line

102

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Islands; BPBM 14788, 1 male, 3 females, Lord Howe
Island; BPBM 15169, 1 male, 1 female, Wake Island, USA;
BPBM 15199, 1 male, 58 mm SL, 6 females, 26-60 mm
SL, Kiritimati, Line Islands; BPBM 15323, 1 male, 4
females, Betio, Tarawa Atoll, Kiribati; BPBM 15616, 1
female, Guadalcanal, Solomon Islands; BPBM 16495, 5
males, 7 females, Pitcairn, Polynesia; BPBM 16534, 2
males, 1 female, Oeno, Pitcairn; BPBM 16607, 1 female,
Oeno, Pitcairn; BPBM 16912, 2 females, Pitcairn; BPBM
16970, 2 males, 31 and 59 mm SL, 5 females, 28-55 mm
SL, Pitcairn; BPBM 16993, 1 female, Pitcairn; BPBM
17042,3 females, Pitcairn; BPBM 17223, 1 male, 1 female,
Mangareva, Gambier Islands, French Polynesia; BPBM
17690, I female, Ryukyu Islands, Japan; BPBM 22357, 2
females, Marshall Islands; BPBM 22834,2 males, 1 female,
Simiian Island, Thailand; BPBM 23262, 1 male and 3
females, Taiwan; BPBM 23338, 2 males and 2 females,
Taiwan; BPBM 33514, 1 male. Chesterfield Bank, Coral
Sea; BPBM 37451, 1 female, Kiritimati; BPBM 38268, 1
female, Tonga; BPBM 38773,2 males, Tahiti; CAS 227288,
43 specimens, Ifalik Atoll, Micronesia; CAS 57943, 3
specimens, Raroia.Tuamotu Islands; CAS 65671, 1 female,
Madang, Papua New Guinea; CAS 81437, 29 specimens,
Raroia, Tuamotu Islands; CAS 227280, 1 juv, 21 mm SL,
24 males 40-63 mm SL, 27 females 42-71 mm SL,
Kapingamarangi Atoll, Micronesia; CAS 81446, 16
specimens, Ifalik Atoll, Micronesia; CAS 81451, 4
specimens, Kapingamarangi Atoll, Micronesia; CAS 81468,
52 specimens, Raroia, Tuamotu Islands; CAS 81483, 96
specimens, Kapingamarangi Atoll, Micronesia; CAS
222525,29 specimens, Fiji;CAS 222527,2 specimens, Fiji;
CAS 222531, 1 female, Fiji; CAS 222537, 1 female, Fiji;
CAS 227283,2 specimens, Fiji; CAS 222549,2 specimens,
Fiji; CAS 222553, 2 specimens, Fiji; CAS 222554, 4
specimens, Yadua Island, Fiji; CAS 222556, 5 specimens,
Fiji; CAS 222569, 1 specimen, Bua Bay, Fiji; CAS 227286,
2 specimens, Vanua Levu, Fiji; CAS 222572, 1 specimen,
Vanua Levu, Fiji; CAS 222582, 5 specimens, Balvu
Harbour, Fiji; CAS 227293, 61 specimens, Fiji; CAS
222601, 45 specimens, Budd Reef, Fiji; CAS 222604, 9
specimens, Fiji; CAS 222606, 20 specimens, Suva, Fiji;
CAS 222607, 40 specimens, Suva, Fiji; CAS 227301, 1
specimen, Suva, Fiji; CAS 222625,8 specimens, Suva, Fiji;
CAS 222626, 13 specimens, Suva, Fiji; CAS 222633, 19
specimens, Suva, Fiji; CAS 222640, 2 specimens, Suva,
Fiji; KAUM-I. 2042, 1 female, 77 mm SL, off Okinoerabu
Island, Kagoshima, Japan, July 1962; KAUM-I. 10652, 1
female, 76 mm SL, off Okinoerabu Island, Kagoshima,
Japan; KAUM-I. 10655, 1 female, 56 mm SL, off
Okinoerabu Island, Kagoshima, Japan; KAUM-I. 10658,
1 male, 49 mm SL, off Okinoerabu Island, Kagoshima,
Japan; KAUM-I. 10663, I male, 33 mm SL, off Okinoerabu
Island, Kagoshima, Japan; KAUM-I. 11482-83, 1 male, 58
mm S Land 1 female, 50 mm SL,30°27'23”N, 130 o 29’59”E,
Isso, Yakushima Island, Kagoshima, Japan; MCZ 158557,
1 specimen, 04°27’S, 171°13’W, Maura Atoll, Kiribati;

MCZ 162572, 4 specimens, 04°27’S, 171°14’W, Manra
Atoll, Kiribati; MCZ 162573, 5 specimens, 04°27’S
171°15’W, Manra Atoll, Kiribati; MNHN 1976-0218, 3
females, 54-56 mm SL, Marshall Islands; MNHN 1980-
0563,2 specimens, New Caledonia; MNHN 1980-0696, 1
female. New Caledonia; MNHN 1980-0961, 1 male, 65
mm SL, 1 female, New Caledonia; NMNZ P.035811, 1
male, 44 mm SL, 1 female, 35 mm SL, Niue, Polynesia;
NSMT-P 55637, 5 specimens, Hainan Island, China;
NSMT-P 55823,2 males, 3 females and 1 juvenile, Hainan
Island, China; ROM 82976, 10 specimens, 18°45’52”S,
178°31 ’13”E, Fiji; ROM 47600,5 specimens, 18°46’30”S,
178°30’28”E, Fiji; ROM 47601,10 specimens, 18°46’13”S,
178°28’05”E, Fiji; ROM 61176,7 specimens, 17 o 31’00”S,
149°55’30”W, Society Islands; ROM 61180, males, 30 and
47 mm SL, female 35 mm SL, 17°36’3r’S, 149°48’42”W,
Society Islands; ROM 82979, I female, 22°21’00”S,
166°2r00”E, New Caledonia; ROM 82978, 1 female, 68
mm SL, 20°48’51 ”N, 107°05’ 13”E, Vietnam; RUSI40747,

7 specimens, Kiritimati, Kiribati; SMNS 17837,1 specimen,
41 mm SL, Aitutaki Island, Cook Islands, 18°50’34”S,

1 59°47’28”W, 0.2-3.5 m depth. SMNS 19762, 11
specimens, 21°38’57”S, 166°18’31”E, New Caledonia,
0-8.5 m; SMNS 25171, 4 specimens, 2I°35’45”S,
167°50'06”E, Loyalty Islands, New Caledonia, 2-3.8 in;
SMNS 21646, 13 specimens, 20°53’35”S, 167°08’02”E,
Loyalty Islands, New Caledonia, 0-2.5 m; SMNS 22732,

1 male, 1 female,20°36’30”S, 164°5r50”E,0-6m; SMNS
22782,1 male, 20 o 3U49”S, 164 o 47’01”E, New Caledonia,
0.2-3.5 m; SMNS 22918, 5 specimens, 20°52’10”S,
167°08’02”E, Loyalty Islands, New Caledonia, 0-3 m;
SMNS 22967, 5 specimens, 20°51’45”S, 167°08’10”E,
Loyalty Islands, New Caledonia, 0-3.5 m; SMNS 23747,

1 male, 1 female, 20°46’54”S, 167°08’19”E, Loyalty
Islands, New Caledonia, 0-3 m; SMNS 23923,8 specimens,
20 o 47 , H”S, 167°07’20”E, Loyalty Islands, New
Caledonia, 0-3 m; SMNS 24849,18 specimens, 21 °56’ 13 ”N,
120°47’53”E, Taiwan; SMNS 24864, 2 specimens,
21°56’13”N, 120°47’53”E, Taiwan; SMNS 24866, 1
specimen, 21°56’13”N, 120°47’53”E, Taiwan; UF 42494,

2 males, 1 female, Enewetak Atoll, Marshall Islands;
USNM 376214, 2 males, 04°0US, 101°0UE, Sumatra,
Mentawai; USNM 384607, 24 specimens, 21°N, 120°E,
Taiwan; USNM 224327,34 specimens, 05°51 ’N, 158°20’E,
Pohnpei, Micronesia, 0-2 m; USNM 297347,59 specimens,
20°24'45”N, 121°55’02”E, Batanes, Philippines, 0-6 m;
USNM 384608, 10 specimens, 18°5UN 121°22’E, Fuga
Island, Philippines; USNM 384604, 17 specimens,
19°51 ’36”S, 174°25’06”W,Tonga, 0-2m; USNM 338463,
18 specimens, 18°40’55”S, 174°06’09”W, Vava’u Group,
Luamoko Island, Tonga, 0-3 m; USNM 384605, 26
specimens, 18 0 44’3r’S, 174°06’36”W, Vava’u Group,
Tonga, 0-11 m; USNM 384593,6 specimens, 20°29’ 12”S,
166°19’18”E, Loyalty Islands, New Caledonia, 3-10 m;
USNM 384601,38 specimens, 19°3r33”S, 169°29’50”E,
Tanna, Vanuatu, I 5 m; USNM 394979, 3 females,

103

W. Schvvarzhans and P. R. Moller

Fig. 10. Alionematichthysphuketensis sp. nov., ZMUC P771659, holotype, male, 40 mm SL.

Fig. 11. Alionematichthys phuketensis sp. nov.. A, lateral view of head, ZMUC ZMUC P771661, male, 37 mm SL; B, ventral view of head,
ZMUC P771661, male, 36 mm SL; C, view of left pseudoclaspers from inside, holotype; D. inclined lateral view of male copulatory organ,
holotype; E, median view of right otolith, holotype; F, ventral view of right otolith, holotype; G, median view of right otolith, ZMUC P771660,
female, 41 mm SL.

13°52’25”S, 167°33’20”E, Banks Islands, Vanuatu, 9-13
m; USNM 366488, 3 females, 28-43 mm SL, Phuket,
Thailand; USNM 366507, 23 males, 40-84 mm SL, 35
females, 39-71 mm SL, Kiriwina Islands (Trobriand),
Papua New Guinea; USNM 366510, 3 males and 19
females, 39-64 mm SL, 05°33’N, 95°09’E, Pulau Boenta,
Sumatra; USNM 366569, 2 females, 62-84 mm SL,
23°25’S, 151°55’E, Heron Island, Queensland, Australia;
USNM 394980, 1 female, 10°35’05”N, I22°08’30”E;
Iloilo, Philippines, 0-7 m; USNM 374192, 1 female, 44
mm SL, 09°06’N, 122°55’E, Negros Island, Philippines;
USNM 376177, 1 male, 45 mm SL, Taiwan; USNM

376178,1 female, 18°56’34”S, 168°59’34”E, Erromango,
Vanuatu, 0-6 m; USNM 376180, 3 males and 7 females,
46-62 mm SL, 08°29’N, 97°39’E, Thailand; USNM
376187, 1 male, 52 mm SL, Sabah; WAM P.29081-016, 3
females, 15°31’S, 123°09’E, Adele Island, Western
Australia, 0.1-2.0 m; WAM P.30633-017, 1 female,
05°09’S, I45°50’E; WAM P.31646-002, 1 male, 13°S
121°E; YCM-P 30026, 1 female, Ishigaki Island, South
Ryukyu Islands, Japan; YCM P 34215, 1 female, Kakeroma
Island, Amami Island, North Ryukyu Islands, Japan.

Additional material. USNM 115392, 1 male, 43 mm
SL, Phoenix Islands, Kiribati; USNM 115395, I male, 52

104

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Fig. 12. Alionematichthyspiger (Alcock, 1890), AMS I.ex. 37315-032, male, 65 mm SL.

mm SL, Phoenix Island, Kiribati; USNM 142010,4 males
and 4 females, 28-42 mm SL, Rongcrik Atoll, Marshall
Islands; USNM 374219,5 males and 5 females, 25—45 mm
SL, Rongelap Atoll, Marshall Islands; USNM 142012, 1
male, 45 mm SL, Kwajalein Atoll, Marshall Islands; USNM
142013, 11 specimens, 40-62 mm SL, Enewetak Atoll,
Marshall Islands; USNM 142014, 9 specimens, 16—40
mm SL, Bikini Atoll, Marshall Islands; USNM 142017,
18 females, 27-58 mm SL, Bikini Atoll, Marshall Islands;
USNM 374220,46 specimens, 32-57 mm SL, Bikini Atoll,
Marshall Islands; USNM 142020, 92 specimens, 28-56
mm SL, Bikini Atoll, Marshall Islands; USNM 374215, 2
males and 1 female, 45-58 mm SL, Amo Atoll, Marshall
Islands; USNM 167355, 3 males and 7 females, 40-55
mm SL, Onotoa, Gilbert Islands, Kiribati; USNM 210154,
1 male, 32 mm SL, Seram, Tandjungliang, Indonesia;
USNM 376161, 86 specimens, 30-84 mm SL, Taiwan;
USNM 263663, 5 males and females, 48-60 mm SL,
Tahiti, French Polynesia; USNM 263674, 19 males and
42 females, 25-56 mm SL, 16°00’S, 175°53’W, Tonga;
USNM 263690, 30 specimens, 29-57 mm SL, 10°55’N,
121°02’E, Palawan, Philippines; USNM 263711, 1 male,
86 mm SL, 21°55’N, 120°49’E, Taiwan; USNM 300090,
1 male and 2 females, 40—45 mm SL, 20°25’N, 121°57’E,
Batanes Islands, Philippines; USNM 300093, 7 males and
8 females, 38-75 mm SL, 20°20’N, 121°49’E, Batanes
Islands, Philippines; USNM 300098, 1 female, 47 mm SL,
20°28’N, 121°58’E, Batanes Islands, Philippines; USNM
300102,1 male and 2 females, 21°07’N, 121°56’E, Batanes
Islands, Philippines; USNM 319899,7 males and females,
45-73 mm SL, Ouvea, Loyalty Islands; USNM 329746, 17
males and 12 females, 31-57 mm SL, 21°18’S, 174°26’W,
Tonga; USNM 333251,3 males and females, 29—41 mm SL,
Tongatapu, Tonga; USNM 336507, 2 females, 55-60 mm
SL, 21°25'S, 174°56’W, Tonga; USNM 374214, 6 males
and 3 females, 35-55 mm SL, 19°16’S, 174°22’W, Tonga;
USNM 338465, 35 specimens, 32-61 mm SL, Vavau,
Tonga; USNM 338966, 7 males and 2 females, 30-45 mm
SL, 18°42’S, 174°02’W, Tonga; USNM 338967, 9 males
and 14 females, 30-50 mm SL, 18°38’S, 174°04'W, Tonga;
USNM 352073, 1 female, 47 mm SL, 16°35’S, 168°06’E,
Lamen Island, Vanuatu; USNM 352742, 123 specimens,

38-80 mm SL, 18°44’S, 169°12’E, Erromango, Vanuatu;
USNM 358502, 18 males and females, 24-71 mm SL,
Tanna, Vanuatu; USNM 361266, 7 females, 35-80 mm
SL, 13°44’S, 167°24’E, Banks Islands, Vanuatu; USNM
376169,1 female, 48 mm SL, 15°00’S, 168°03’E, Maewo,
Vanuatu; USNM 366200, 5 males and females, 17-65
mm SL, Tonga; USNM 366222, 5 males and females,
38-61 mm SL, Line Islands; USNM 366223, 1 male, 51
mm SL, Starbuck Atoll, Line Islands; USNM 366225,
1 male, 2 females, Bora Bora, Society Islands; USNM
366489, 2 females, 34-49 mm SL, Taka Atoll, Marshall
Islands; USNM 376218, 1 female, 44 mm SL, 04°14’S,
152°10’E, Bismarck Archipelago, Papua New Guinea;
USNM 366511, 2 females, 36 mm SL, Amami, Kikai
Island, Ryukyu Islands; USNM 376213, 2 females, 38-63
mm SL, Kiriwina Islands (Trobriand); USNM 366556, 1
female, 38 mm SL, Taiwan; USNM 366560, 1 female, 49
mm SL, 25°12’N, 121°4UE, Taiwan; USNM 366561, 12
specimens, 43-78 mm SL, 25°12’N, 121 °41 ’E, Taiwan;
USNM 366574, 2 males and 8 females, 44-51 mm SL,
Kwajalein Atoll, Marshall Islands; USNM 366575, 1 male,
35 mm SL, Phoenix Atoll, McKean Island; USNM 374221,
1 female, 52 mm SL, Kwajalein Atoll, Marshall Islands;
USNM 366719, 1 male and 2 females, 51-67 mm SL, Jarvis
Island, Line Islands; USNM 366725,2 males and 1 female,
40-51 mm SL, Bora Bora, Society Islands; USNM 366843,
1 male and 1 female, 54-55 mm SL, 20°37’S, 178°40’E,
Fiji; USNM 376211, 1 male and 4 females, 30-48 mm SL,
18°57’S, 178°17’E, Fiji; USNM 366847, 1 male, 49 mm
SL, 18°56’S, 178°21 ’W, Fiji; USNM 366848,2 males and
4 females, 32-45 mm SL, 17°47’S, 177°13'E, Fiji.

Diagnosis. Vertebrae 11-12+29-33=40-45, dorsal fin
rays 72-88, anal fin rays 55-70; eyes moderately large (1.7-
2.8% SL); body slender, snout rounded, with many cirri;
scales only on cheeks (rarely 1-2 scales above opercular
spine); upper preopercular pore present; outer pseudoclasper
broad-based, large, often extending beyond hood in resting
position; inner pseudoclasper small, anterior and posterior
lobes of equal size; otolith length to height 1.9-2.1, with
gently curved dorsal rim, otolith length to sulcus length
1.5-1.6, ostium length to cauda length 3.5-4.5.

105

W. Schwarzhans and P. R. Moller

Fig. 13 (part). Alionematichthys piger (Alcock, 1890), A, lateral view of head, ROM 82976, male, 5 1 mm SL; B, lateral view of head, USNM
384707, male, 61 mm SL; C, lateral view of head, USNM 366507, male, 54 mm SL; 1), ventral view of head, USNM 384707, male, 61 mm
SL; E, lateral view of head, CAS 227288, female, 51 mm SL; F, inclined lateral view of male copulatory organ, USNM 376 1 80, male, 57 mm
SL; G, view ot left pseudoclaspers from inside, USNM 376180, male, 57 mm SL; H, view of left pseudoclaspers from outside, USNM 376180,
male, 44 mm SL; !, ventral view of male copulatory organ, USNM 366507, male, 54 mm SL; J, view oflcft pseudoclaspers from inside,
SMNS 1 7837,41 mm SL; K, view of left pseudoclaspers from inside, ROM 82976,43 mm SL; L, view of left pseudoclaspers from inside,
MNI IN 1980-0961,65 mm SL; M, view of left pseudoclaspers from inside, CAS 227288, male, 59 mm SL; N, view of left pseudoclaspers
from inside, BPBM 8009, male, 43 mm SL; O, ventral view of female copulatory organ, SMNS 19762, 85 mm SL; P. inclined lateral view
of female copulatory organ, SMNS 19762, 85 mm SL.

106

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Fig. 13 (continued). Q, median view of right otolith, USNM 297347, male, 67 mm SL; R, ventral view of right otolith, USNM 297347, male,
67 mm SL; S. median view of right otolith, USNM 384605, male, 52 mm SL; T, median view of right otolith, AMS 1.23381-001, isolated
otolith; U, median view of right otolith, MNHN 1976-0218, female, 56 mm SL.

Description (Figs 12, 13). The principal meristic and
morphometric characters are shown in Table 5. Mature at
about 40 mm SL. Body slender, with rounded head profile.
Many cirri on snout. Eye size 1.7-2.8 (2.6)% SL. Head
with scale patch on cheek containing 6-9 vertical rows of
scales on upper part and 3^1 vertical rows on lower part.
Horizontal diameter of scales on body of a 65 mm SL male
(AMS 1.37315-032) about 1.8% SL, in about 27 horizontal
rows. Maxillary ending far behind eye, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior end
expanded. Anterior nostril positioned high, 1/2.5 distance
from upper lip to anterior margin of eye. Posterior nostril
small, about one-fifth to one-sixth the size of eye. Opercular
spine with free tip, thin and sharply pointed. Anterior gill
arch with 13-19 rakers, 3 elongate, sometimes with short
plate-like raker between the two lower elongate rakers.
Pseudobranchial filaments 2.

Head sensory’pores (Fig. 13A-E). Supraorbital pores 2
to 3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about 1/3 the size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior): first anterior
pore large, tubular, without cirri. Preopercularpores: 3 lower,
first and second with joined opening, covered by dermal flap
in lateral view; third nontubular; upper preopercular pore
present. [See description of Alionematichthys ceylonensis
for position of pores.]

Dentition (of a 59 mm SL specimen - USNM 376180).
Premaxilla with 5 outer rows of granular teeth and 2 inner

row of larger teeth anteriorly. Anteriormosl teeth in inner
row up to 1/2 diameter of pupil. Vomer horseshoe-shaped,
with 2 outer rows of small teeth and I inner row of larger
teeth up to 1/3 diameter of pupil. Palatine with 2 outer
rows of small teeth and 1 inner row of larger teeth up to 1/3
diameter of pupil. Dentary with 5 outer rows of granular
teeth and 1 inner row of larger teeth anteriorly, up to about
3/4 diameter of pupil.

Otolith (Fig. 13Q-U). Moderately elongate in shape,
length to height 1.9-2.1 and thin (otolith height to otolith
thickness about 2.3). Anterior tip broadly rounded, posterior
tip expanded and irregularly ornamented. Dorsal rim gently
curved, without distinct angles. Inner face moderately
convex, outer face almost flat, both smooth. Otolith length
to sulcus length 1.5-1.6. Sulcus slightly supramedially
positioned, slightly inclined, with separated colliculi and
marked notch at ventral rim of sulcus at joint of ostium
with cauda. Ostium length to cauda length 3.5-4.5. Ventral
furrow very distinct and close to ventral rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined and
5-8 depressed. Parapophyses present from vertebrae 6 to
II (12). Pleural ribs on vertebrae 2 to 10(11). First anal
fin plcrygiophore not or slightly longer than subsequent
pterygiophore.

Male copulatory organ (Fig. 13G-N). Two pairs of
pseudoclaspers, outer usually very large and extending
beyond hood in resting position. Outer pseudoclasper a
large, flat flap with broad base; inner pseudoclasper short,

107

W. Schwarzhans and P. R. Moller

sometimes compressed, with anterior thorn and posterior
lobe being equal in size. Isthmus narrow; penis curved,
about as long as outer pseudoclaspers.

Female copulatory organ (Fig. 130, P). Few female
specimens of A. piger show soft dermal flaps around the
vagina which may reach a quite considerable size, as in the
figured specimens but do not exhibit diagnostic valuable
morphology.

Colour. According to Alcock (1890:433), the live colour
is uniform dark brown, almost black. The preserved colour
is similar.

Variability. Alionematichthyspiger is remarkable for an
unusual wide variation in pseudoclasper morphology that
does not follow a geographic pattern. The majority of the
male specimens show a relatively large outer pseudoclasper
for a member of this genus, extending beyond the hood
in resting position and about twice as long as the inner
pseudoclasper. In extreme forms the inner pseudoclasper
is depressed (Fig. 13J, K.). On the other extreme, the inner
pseudoclasper is rather stubby and the outer pseudoclasper
less than half the length of the inner pseudoclasper and not
extruding in resting position (Fig. 13M, N). The very large
variation is also reflected in the meristic counts (Tables
1, 5) and variations in the amount of cirri on the snout
(Fig. 13A-E) which may represent geographic variances.
Otherwise, heads (Fig. 13A-E) and otoliths (Fig. 13Q-U)
of fishes with such diverse pseudoclaspers do not document
matching variation.

Remarks. Alcock (1890) described Dinematichthys
piger from Great Coco Island, Andaman Islands. The
unique holotype of D. piger (ZSI F12939) was not available
for review (see also Schwarzhans and Moller 2007: 66).
According to Alcock’s description it is a 61 mm long
specimen with 75 dorsal fin rays, 55 anal fin rays and 90
scales along the lateral line and his figure shows a fish with a
rather high position of the anterior nostril at 1 /3 the distance
from upper lip to aggregate distance to anterior margin of
eye and no scales on the opercle.

The latter two observations agree with a species of
the genus Alionematichthys. There are two species of the
genus Alionematichthys observed along the shores of the
Andaman Sea, one without scales above the opercle, the
other with many scales above the opercle. The latter has
been attributed to a species described from the Ryukyu
Islands, namely Dinematichthys riukiuensis, now placed in
the genus Alionematichthys (see below). Another species
described above from the Thailand coast, A. phuketensis,
across the Andaman Sea from Alcock’s type locality, does
not seem to attain such size and has higher dorsal (81-89)
and anal (60-65) fin ray counts. Finally, Alcock’s drawing
is detailed enough to reveal that the holotype must be a
female. With all these facts in mind it has to be considered
that the specimens attributed to A. piger in the following
and the diagnosis of the species and description are not fully
satisfactory until the holotype has been re-studied.

Comparison. Alionematichthys piger belongs to the
species group with an upper preopercular pore, large eyes
and no (rarely 1 or 2) scales above the opercular spine
comprising also A. samoaensis sp. nov. and A. suluensis
sp. nov., which both have a much more restricted distribution.
It differs from A. samoaensis sp. nov. by the lack of a knob
at the distal inner end of the outer pseudoclasper and the
simple flap at the anterior margin of the posterior nostril
(vs strongly elevated funnel shaped). From A. suluensis, it
differs in the anterior lobe of the inner pseudoclasper being
equally long as its posterior lobe (vs very long anterior
lobe, twice as long as posterior lobe) and many cirri on the
snout (vs no cirri).

Distribution. Alionematichthys piger is one of the most
common and most widely distributed Dinematichthyini
(Fig. 9). Its distribution ranges from the Andaman Islands
to Sumatra, Thailand, Vietnam, Taiwan and the Ryukyu

Table 6. Meristic and morphometric characters of Alionematichthys
piger ( Alcock, 1890)

Holotype

Holotype +
340 non-types

Mean (range)

Standard length in mm

Meristic characters

50.6 (26-86)

Dorsal fin rays

79.7 (72-88)

305

Caudal fin rays

16.0(14-18)

Anal fin rays

61.8(55-70)

305

Pectoral fin rays

20.6(19-22)

Precaudal vertebrae

11.1 (11-12)

340

Caudal vertebrae

31.1 (29-34)

337

Total vertebrae

42.1 (40-45)

337

Rakers on anterior gill arch

15.8(13-19)

Pseudobranchial filaments

D/V

5.7(5-7)

338

D/A

22.7 (19-28)

336

V/A

Morphometric characters in % of SL

13.0(12-16)

337

Head length

25.1

25.7(24.0-27.5)

Head width

14.0(11.8-15.0)

Head height

16.7

15.9(13.6-16.8)

Snout length

5.8(54-6.6)

Upper jaw length

13.5 (12.5-14.4)

Diameter of pigmented eye

2.6

2.3(1.7-2.8)

Diameter of pupil

1.4 (0.9-1.6)

Interorbital width

7.1 (6.5-7.9)

Posterior maxilla height

4.5 (4.0-5.2)

Postorbital length

17.8

18.6(17.6-19.5)

Preanal length

50.4

48.0 (44.4-50.9)

Predorsal length

31.2

30.1 (27.4-31.2)

Body depth at origin of anal fin

16.6

17.8(14.8-19.5)

Pectoral fin length

13.9

13.5(11.9-14.7)

Pectoral fin base height

■ 5.7

6.1 (5.7-6.9)

Ventral fin length

24.0(21.2-27.6)

Base ventral fin - anal fin origin

29.7(25.3-34.5)

* Data taken from Alcock (1890 and 1905).

108

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Islands in the north, to Australia and as far south as Lord
Howe Island and eastwards to Micronesia, the Line Islands,
the Tuamotu Archipelago and Pitcairn Group. Together with
Diancistrus katrineae Schwarzhans, Moller and Nielsen, it
represents the species farthest to the east in the West Pacific
at Ducie Atoll. The species is found in holes and crevices
of coralline and volcanic rock.

Alionematichthysplicatosurculus sp. nov.

(Figs 1, 14, 15, Tables 1,7)

Material examined. (152 specimens, 28-79 mm
SL). HOLOTYPE - USNM 384198, male, 79 mm SL,
10°34’45”N, 122°30’30”E, 0-4 m, small cove lined with
volcanic rock and with rocky boulders in cove, volcanic
rock boulders and sandy channels, coral over part of
area; Guimaras Island, Pulang Duta, Sinabsapan, Panay,
Philippines, J. T. Williams, D. G. Smith, K. E. Carpenter,
C. Carpenter, N. Minsalan and T. Minsalan, 24 Sept.
1995. PARATYPES - USNM 209801. 1 male, 53 mm SL,
Indonesia, Ambon, off Tandjung Suli, shallow coral reef,
0-2 m, V. G. Springer and M. F. Gomon, II Jan. 1973;
CAS 227311, 1 female, 45 mm SL, 05°14’S, 145°47’E,
Papua New Guinea, Madang, Nagada Harbour, 1-7 m,
S. G. Poss, D. Catania et al., 5 December 1987; USNM
365822, 2 males, 57 and 62 mm SL, Papua New Guinea,
Papua, Harvey Bay, east of Oro Bay, 0-10 m, T. R. Roberts,
6-7 August 1975; USNM 365832, 1 male, 55 mm SL, 2
females, 46 and 58 mm SL, 05°11’S, I45°50’E, Papua
New Guinea, Kranket Island, mangrove right up to coral,
0-1 m, V. G. Springer etal., 7 Nov. 1978; USNM 366567,
1 male, 50 mm SL, Solomon Islands, Bougainville, Kieta
Harbor, coral reef offmain market, 1-4 m, D. M. Cohen and
party, 10 Mar. 1965; USNM 366596, 1 female, 44 mm SL,
09°06’30”N, 122°55’24'’E, Philippines, ncarGiligaon, N
ofMaloh, Negros, 0-2 m, L. W. Knapp etal. 26 April 1979;
USNM 376183, I male, 47 mm SL, 04°14'S, 152°10’E,
Papua New Guinea, Bismarck Archipelago, Rabaul, New
Britain, W. Davis et al., 27 Feb. 1965; USNM 394981, 3
females, 40, 52 and 62 mm SL, 1 male 48 mm SL, same
data as holotype; WAM P.30635-001, 1 male, 45 mm SL,
Papua New Guinea. Madang, 0-4 m, G. R. Allen et al., 10
Feb. 1993; WAM P.29624-044, 5 females, 37-52 mm SL,

Table 7. Meristic and morphometric characters of Alionematichthys
plicatosurculus sp. nov.

Holotype

Holotype +

29 paratypes

USNM

384198

Mean (range)

Standard length in mm

Meristic characters

48.9 (28-79)

Dorsal fin rays

80.3 (76-84)

Caudal fin rays

Anal fin rays

62.6 (60-67)

Pectoral fin rays

20.2(19-21)

Precaudal vertebrae

12.0(11-12)

Caudal vertebrae

31.1 (30-33)

Total vertebrae

43.1 (42-45)

Rakers on anterior gill arch

16.3(14-18)

Pseudobranchial filaments

D/V

5.9 (5-6)

D/A

22.3 (19.0-25.0)

VIA

13.4(11.0-19.0)

Morphometric characters in % of SL

Head length

24.7

26.0(24.7-28.1)

Head width

14.5

13.6(11.7-15.6)

Head height

16.6

16.0(13.8-17.7)

Snout length

5.3

5.9(5.1-7.3)

Upper jaw length

13.3

13.4(12.5-14.5)

Diameter of pigmented eye

2.2

2.6 (2.2-3.0)

Diameter of pupil

1.2

1.6(1.2-2.2)

Interorbital width

7.5

7.2 (6.7-8.1)

Posterior maxilla height

4.5

4.4 (3.8-4.8)

Postorbital length

17.8

18.5(17.6-19.2)

Preanal length

45.9

46.1 (43.5-47.7)

Predorsal length

30.4

31.3 (29.8-33.0)

Body depth at origin of anal fin

18.2

18.4(17.1-19.4)

Pectoral fin length

13.6

13.9(11.9-16.0)

Pectoral fin base height

6.0

5.9 (5.2-6.7)

Ventral fin length

22.0

23.5(19.4-25.8)

Base ventral fin - anal fin origin

22.3

26.6 (22.3-28.5)

9 males, 28-52 mm SL, 05°09’S, 145°48’E, Papua New
Guinea, Madang, G. R. Allen, 15 Oct. 1987.

Fig. 14. Alionematichthys plicatosurculus sp. nov., USNM 394198, holotype, male, 79 mm SL.

109

W. Schwarzhans and P. R. Moller

Fig. 15. Alionemalichthys plicatosurcttlus sp. nov.. A, lateral view of head, holotype; B. ventral view of head, holotype; C, lateral view of
head, paratypc, USNM 394981, female, 52 mm SL; D, inclined lateral view ofmalecopulatory organ, holotype; E, view of left pscudoclaspcrs
from inside, holotype; F, ventral view of left pseudoclaspers, holotype; G, ventral view of male copulatory organ, holotype; H, view of left
pseudoclaspers from inside, USNM 366567, 50 mm SL; I, median view of right otolith, holotype; J, ventral view of right otolith, holotype.

110

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Additional material. BMNH 1907.2.28.11,1 specimen,
(bad condition), Neu-Lauenburg/ Duke of York Islands,
Bismarck Archipelago, Papua New Guinea; USNM
263681,33 males and 52 females, 30-70 mm SL, 09°06’N,

122°55’E, Philippines, Negros; USNM 263683,4 males and
9 females, 33-71 mmSL, 09°03’N, 122°59’E, Philippines,
Negros; USNM 377193, 1 male and 3 females, 68-72
mm SL, 09°10’N, 123°26’E, Philippines, Siquijor Island;
USNM 377202, 8 specimens, 48-71 mm SL, 10°35’N,
122°08’E, Philippines, Panay Island; USNM 345618, 5
specimens, 11°25’N, 123°15’E, Panay, Philippines; USNM
365818,20 specimens, Papua New Guinea, Bagabag Island;
USNM 366480, 1 female, 52 mm SL, 05°14’S, I45°47’E,
Papua New Guinea, Madang; USNM 3766222, 1 male, 66
mm SL, 05°14'S, 145°47’E, Papua New Guinea, Madang,
USNM 366508, 3 females, 40-60 mm SL, 03°23’S,
I43°40’E, Papua New Guinea, Mushu Island.

Diagnosis. Vertebrae 11-12+31-33=42-45, dorsal fin
rays 76-84, anal fin rays 60-67; eyes moderately large
(2.2—3.0% SL); body slender, snout rounded, without cirri;
scales only on cheeks; no upper prcopercular pore; outer
pseudoclasper broad-based, large, sometimes extending
beyond hood in resting position, with thickened tip,
particularly on the inner face; inner pseudoclasper large,
thick, with posterior lobe folded over anterior pointed lobe;
otolith length to height 2.1 -2.3, with gently curved dorsal
rim, otolith length to sulcus length 1.6-1.7. ostium length
to cauda length 3.5=4.5.

Description (Figs 14-15).The principal meristic and
morphometric characters are shown in Table 7. Mature at
about 40 mm SL. Body slender, with rounded head profile.
No cirri on snout.

Eye size 2.2-3.0 (2.2)% SL. Head with scale patch
on cheek containing 5-6 (6) vertical rows of scales on
upper part and 2-3 vertical rows on lower part. Horizontal
diameter of scales on body of holotype about 1.7% SL,
in 32 horizontal rows. Maxillary ending far behind eye,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded, with small knob. Anterior
nostril positioned high, 1/2.5—1/3 distance from upper lip
to anterior margin of eye. Posterior nostril small, about 1/5
to 1/6 the size of eye. Opercular spine with free tip, thin and
sharply pointed. Anterior gill arch with 14-18(15) rakers,
3 elongate in row. Pseudobranchial filaments 2.

Head sensory pores (Fig. 15 A-C). Supraorbital
pores 3. Infraorbital pores 6 (3 anterior and 3 posterior),
three posterior pores about one-quarter the size of three
anterior pores. Mandibular pores 6 (3 anterior and 3
posterior); first anterior pore large, tubular, without cirri.
Prcopercular pores: 3 lower, first and second with joined
opening, covered by dermal flap in lateral view; third
tubular; upper preopercular pore absent. [See description of
Alionematichthys ceylonensis for position of pores.]

Dentition (of holotype). Premaxilla with 7 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 1/4 diameter of pupil.

Vomer horseshoe-shaped, with 2 outer row of small teeth
and 1 inner row of large teeth up to 1/4 diameter of pupil.
Palatine with 3 outer rows of small teeth and 1 inner row
of long teeth up to 1/3 diameter of pupil. Dentary with 5
outer rows of granular teeth and 1 inner row of larger teeth
anteriorly, up to about 1/2 diameter of pupil.

Otolith (Fig. 151, J). Moderately elongate in shape,
length to height 2.1-2.3 and thin (otolith height to otolith
thickness about 3). Anterior tip broadly rounded to
slightly angular, posterior tip much expanded and slightly
ornamented. Dorsal rim gently curved, without distinct
angles. Inner face moderately convex, outer face concave
to almost flat, both smooth. Otolith length to sulcus length
1.6-1.7. Sulcus slightly supramedially positioned, slightly
inclined, with separated colliculi and marked notch at
ventral rim of sulcus at joint of ostium with cauda. Ostium
length to cauda length 3.5-4.5. Ventral furrow feeble and
close to ventral rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined
and 5-8 depressed. Parapophyses present from vertebrae
6 to 11 (12). Pleural ribs on vertebrae 2 to 10(11). First
anal fin pterygiophore slightly longer than subsequent
pterygiophore.

Male copulatory organ (Fig. 15D-H). Two pairs of
moderately large pseudoclaspers. Outer pseudoclasper large,
sometimes extending beyond hood in resting position, with
thickened tip (particularly in large specimens) expressed
as massive knob on inner face; inner pseudoclasper large,
thick, with posterior lobe very' large and folded over anterior
pointed lobe. Isthmus moderately narrow; penis curved,
about as long as outer pseudoclaspers.

Colour. Live colour unknown. Preserved colour
uniformly brownish.

Comparison. Alionematichthys plicatosurculus
belongs to the species group without an upper preopercular
pore together with A. ceylonensis, A. phuketensis and
A. shinoharai sp. nov., from which it differs in the peculiar
shape of the thick inner pseudoclasper with the posterior
lobe folded over the anterior lobe and the massive tip of the
inner face of the outer pseudoclasper. For further distinction
from A. ceylonensis and A. phuketensis sec respective
descriptions. From A. shinoharai sp. nov. it differs further in
the lack of scales above the opercular spine (vs 2), the long
sulcus (otolith length to sulcus length 1.5-1.6 vs 1.9-2.0)
and the notch at the ventral sulcus margin at the joint of
ostium and cauda (vs no notch).

Distribution. Alionematichthys plicatosurculus is
known from the Philippines south of 12°N, Ambon and
along the northern coast of New Guinea to the Bismarck
Archipelago and Solomon Islands (Fig. 1).

Etymology. Named after the characteristic folded inner
pseudoclasper by a combination ofplicatus (Latin = folded)
and surcutus (Latin = grapevine tendril) used for the term
pseudoclasper. It is a noun in apposition.

W. Schvvarzhans and P. R. Mollcr

Fig. 16. Sample sites of O Alionematichthys riukiuensis (Aoyagi, 1954), * Alionematichthys aff. riukiuensis 1. * Alionematichthys aff.
riukiuensis 2, © A. samoaensis sp. nov., © A. winlerbottomi sp. nov., © Alionematichthys sp. 1, © Alionematichthys sp. 2. One symbol may
represent several samples.

Fig. 17. Alionematichthys riukiuensis (Aoyagi, 1954), A, fresh dead, WAM P.30909-002 [exact specimen and size not known],
B, NMST-P 55637, male,74 mm SL.

112

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Alionematichthys riukiuensis (Aoyagi, 1954)

(Figs 16-19, Tables 1,8)

Dinematichthys riukiuensis Aoyagi, 1954: 235;
Machida 1992: 270; Machida 1994: 461; Hayashi 1995:
11; Nielsen et al. 1999: 130.

Brotulinafusca. — Matsubara 1955:800 (not Diancistrus
fuscus Fowler, 1946).

Dinematichthys megasoma Machida, 1994:459; Nielsen
et al. 1999: 130; Schwarzhans and Mollcr2005: 78.

Material examined. 536 specimens, 19-150 mm SL.
PARALECTOTYPE - YCM-P 30001, female, 72 mm,
Okinawa Island, Ryukyu Islands; NON-TYPES - AMS
1.37399-076, male, 73 mm SL, Vanuatu; AMS 1.18739-037,
1 male and 2 females, 14°42’S, 145°27’E, Lizard Island,
Queensland, Australia; AMS 1.19108-032, 2 males and 3
females, 14°40’S, I45°28’E, Lizard Island, Great Barrier

Table 8. Meristic and morphometric characters of Alionematichthys
riukiuensis (after Aoyagi 1954).

Holotype

YCM-P

30023*

Holotype +
non-types

Mean (range)

Standard length in mm

Meristic characters

72.3(20-148)

121

Dorsal fin rays

83.9 (77-92)

107

Caudal fin rays

15.7(14-16)

Anal fin rays

65.1 (59-72)

107

Pectoral fin rays

20.5(19-23)

Precaudal vertebrae

11.3(11-12)

121

Caudal vertebrae

31.5(29-33)

120

Total vertebrae

42.8 (41-44)

120

Rakers on anterior gill arch

19.1 (15-21)

Pseudobranchial filaments

1.9(0-3)

D/V

5.7 (5-7)

111

D/A

23.7(19-27)

112

VIA

Morphometric characters in % of SL

13.1 (12-14)

112

Head length

29.3

26.8 (25.7-29.3)

Head width

14.8(11.6-18.6)

Head height

17.1 (14.3-20.5)

Snout length

7.3

5.8 (4.7-7.3)

Upper jaw length

14.3

13.9(12.4-15.1)

Diameter of pigmented eye

3.0

2.6 (2.5-3.0)

Diameter of pupil

1.4 (1.2-1.6)

Interorbital width

7.7

7.3 (6.2-8.0)

Posterior maxilla height

5.3

4.6(3.8-5.3)

Postorbital length

20.6

19.5(19.0-20.6)

Prcanal length

49.8

47.1 (45.7-49.8)

Predorsal length

33.0

31.2 (29.9-33.0)

Body depth at origin of anal fin

21.4

19.9(16.8-23.4)

Pectoral fin length

15.7

14.4(12.8-16.1)

Pectoral fin base height

7.5

6.1 (4.4-7.5)

Ventral fin length

20.4

21.6(20.0-24.8)

Base ventral fin - anal fin origin

26.5(21.8-30.9)

* Data from Machida (1994) included.

Reef, Australia, 1-10 in, D. F. Hoese and party, 17 Nov.
1975; AMS 1.20770-015,42 specimens, 11 °55’S, 143°27’E,
Sir Charles Hardy Island, Queensland, Australia; AMS
1.20770-073,11 specimens, 11 °55’S, 143°27’E, Sir Charles
Hardy Island, Queensland, Australia; AMS 1.21422-029,5
specimens, 14°S, 145°E, Coral Sea; AMS 1.33740-026, 1
male, 10°S 144°E, Coral Sea; AMS 1.34311-010, 1 male,

5 females, 22°14’S, 150°19’E, Australia, Queensland,
Cannibal Group; AMS 1.37315-032,3 males and 4 females,
18°44’S, 169°12’E, Vanuatu, Erromango; AMS 1.37322-
009,4 females, 19°31’S, 169°29’E, Vanuatu, Tanna; AMS
1.37339-076, 1 male and 1 female, 16°47’S, 168°21 ’E,
Vanuatu, Epi; AMS 1.37922-021, 1 male, and 1 female,
13°52’S, 167°32’E, Vanuatu. Banks Islands; BPBM 5921,

1 female, Vanuatu, Efate; BPBM 11465, 2 females, New
Caledonia; BPBM 40934, 1 male, 2 females, Ryukyu
Islands; BPBM 40936,2 females, Similan Island Thailand;
CAS 65664, I male, Papua New Guinea, Madang; KAUM-I.
11459,1 male, 61 mmSL,30°27’23”N, 130°29’59”E, Isso,
Yakushima Island, Kagoshima, Japan; KAUM-I. 10661,

1 male, 48 mm SL, off Okinoerabu Island, Kagoshima,
Japan; KAUM-I. 11484, 1 male, 55 mm SL. 30°27’23”N,
130°29’59”E, Isso, Yakushima Island, Kagoshima, Japan;
KSHS 3650 (otolith only); MNHN 1980-0851, 3 females.
New Caledonia; NSMT-P 55636, female 74 mm SL,
China, Hainan Island, China; NSMT-P 55637, 1 male,
74 mm SL, Hainan Island, China; NTM S. 13425-007, 1
female, 11°58’S, 123°2 HE, Ashmore Islands, Australia;
ROM 71841, 9 specimens, Vietnam; ROM 71843, 11
specimens, Vietnam; ROM 71850, 5 specimens, 20°N
107°E; SMNS 26429, 5 specimens, eastsoutheast ofThio,
New Caledonia, 21°38’57”S, 166°18’31”E, 0-8.5 m;
SMNS 26430, 6 specimens, Mare, Cap Wabao, Loyalty
Islands, 21°35’45 ,, S 167°50’06”E, 2-3.8 m; SMNS 22705,

6 specimens, northnorthwest of Ouegoa, New Caledonia,
20°15’20”S, 164°24’10”E, 0.5-3.5 m; SMNS 22804, 5
specimens, east-south-east of Ponerihuen, New Caledonia,
21°04’55”S 165°28'10”E, 0.3-2.8 m; SMNS 26431, 2
specimens, north-west of Hienghene, New Caledonia,
20°31’49”S 164°47’0r’E, 0.2-3.5 m; USNM 199541, 9
males and 14 females, 42-66 mm SL, 04°01’S, 101 °01 ’E,
Mentawai, Sumatra; USNM 394978, 5 specimens, 21°N
120°E; EX USNM 345618, 7 males, 29-65 mm SL, 1
female, 40 mm SL, juvenile 20 mm SL, 11°25’N, 123°15’E,
Panay, Philippines; USNM 394982,31 specimens, 16°47’S,
168°2UE, Epiu, Vanuatu; USNM 355385, 38 specimens,
Tanna, Vanuau; USNM 394984, 1 female, 75 mm SL,
17°31’S, 168°19\ Efate, Vanuatu; USNM 362434, 1
male, 64 ram SL, 3 females, 4 juveniles, 14°S 167°E,
Banks Islands, Vanuatu; USNM 366490, 3 males, 72-78
mm SL, 3 females, 57-72 mm SL, 08°29’N, 97°39’E,
Similan Islands, Thailand; USNM 366600, 1 male, 62
mm SL, 09°19’N, 123° 18’E, Negros, Philippines; USNM
376188, 1 male, 150 mm SL, 2 females, 116-119 mm SL,
Taiwan; WAM P.30305-037, 1 male, 2 females, 13°52’S,
126°56’E, Sir Graham Moore Islands, Western Australia;

113

W. Schwarzhans and P. R. Moller

Fig. 18. Alionematichthys riukiuensis (Aoyagi, 1954), A, lateral view of head, ROM 71841, male, 73 mm SL; B, lateral view of head, WAM
P.30308-001, male. 112 mm SL; C, ventral view of head, WAM P.30308-001, male, 112 mm SL; D, laieral view of head, AMS 1.34311-
010, female, 69 mm SL; E, inclined lateral view of male copulatory organ, USNM 394982, 92 mm SL; F, view of left pseudoclaspers from
inside, USNM 394982, 92 mm SL; G, ventral view of male copulatory organ, SMNS 26430, 130 mm SL; H, view of left pseudoclaspers
from inside, USNM 376188, 150 mm SL; I. ventral view of left pseudoclaspers, WAM P.30308-001, 112 mm SL; J, median view of right
otolith, WAM P.30308-001, 112 mm SL.

114

Dinematichthyine fishes of the Indo-west Pacific, Part IV

WAM P.30308-001,7 specimens, 13°S, 125°E, Timor Sea;
WAM P.30909-002,8 specimens, 16°S, 123°E, North West
Australia; YCM-P44101, 1 male and 2 females, Ishigaki,
Ryukyu Islands; YCM-P44102, 3 specimens, Kakeroma
Island, Ryukyu Islands.

Additional material. USNM 151418,1 male, 59mm SL,
Philippines; USNM 151420,1 male, 74 mm SL, Philippines;
USNM 151422, 1 female 73 mm SL, Philippines; USNM
151423, 1 female, 82 mm SL, Philippines; USNM 151424,

1 male, 84 mm SL, Philippines; USNM 151425, 1 female,
102 mm SL, Philippines; USNM 151426, 1 female, 50
mm SL, Philippines; USNM 164589, 1 female, 78 mm
SL, Espiritu Santo, Vanuatu; USNM 376160, 1 male and
3 females, 62-70 mm SL, Taiwan; USNM 377258, 15
specimens, 38-83 mm SL, 02°35’S, 150°47’E, Kavieng,
New Ireland, Papua New Guinea; USNM 263666, 5 males
and 9 females, 34-83 mm SL, 16°25’N, 119°54’E, Luzon,
Philippines; USNM 263680, 2 males, 59-109 mm SL,
23°29’S, 152°05’E, One Tree Island, Queensland, Australia;
USNM 377214, 25 specimens, 47-109 mm SL, 09°06’N,
122°55’E, Negros, Philippines; USNM 376347, I female,
61 mm SL, 10°52’N, 120°56’E, Palawan, Philippines;
USNM 376225, 1 female, 49 mmSL, 10°55’N, 121°02’E,
Palawan, Philippines; USNM 263700, 1 male, 68 mm
SL, 09°04’N, 123°16’E, Apo Island, Philippines; USNM
263743, 5 specimens, 34-76 mm SL, Milne Bay, Papua
New Guinea; USNM 263755, 15 specimens, 31-119 mm
SL, Taiwan; USNM 377208, 1 male and 3 females, 79-104
mm SL, 10°35’N, 122°08’E, Panay, Philippines; USNM
376163, 6 females, 68-92 mm SL, 18°44’S, 169°12’E,
Erromango, Vanuatu; USNM 355152, 1 female, 100 mm
SL, 17°03’S, 168°21’E, Emae, Vanuatu; USNM 356612,

10 specimens, 19-95 mm SL, Tomotu, Santa Cruz Islands,
Solomon Islands; USNM 376167, 3 females, 35-51 mm
SL, 13°32’S, 167°20\ Banks Islands, Vanuatu; USNM
363192, 1 male and 1 female, 33-75 mm SL, Banks
Islands, Vanuatu; USNM 376166, 3 females, 30-88 mm
SL, 13°04’S, 167°39’E, Banks Islands, Vanuatu; USNM
365819,7 specimens, 44-72 mm SL, Bagabag Island, Papua
New Guinea; USNM 376223,2 males and 4 females, 55-80
mm SL, 05°14'S, I45°47’E, Madang, Papua New Guinea;
USNM 366482, 16 males and 39 females, 32-75 mm SL,
10°47’S, 152°24’E, Louisade Archipelago, Deboyne Atoll,
Papua New Guinea; USNM 376202, 1 female, 55 mm SL,
05°14’S, 145°47’E, Madang, Papua New Guinea; USNM
376190,1 female, 65 mmSL,05°14’S, 145°47’E, Madang,
Papua New Guinea; USNM 366521,10 specimens, 32-80
mm SL, 05°10’S, 145°51’E, Kranket Island, Papua New
Guinea; USNM 366522,2 males and I female, 40 mm SL,
01°31’S, 145°01 ’E, Hermit Islands, Papua New Guinea;
USNM 366524, 1 male, 67 mm SL, 05°14’S, 145°47’E,
Madang, Papua New Guinea; USNM 366557, 1 male and
1 female, 51-73 mm SL, 21°55’N, 120°44’E, Taiwan;
USNM 366559, 1 female, 56 mm SL, Taiwan; USNM
374226, 6 specimens, 35-103 mm SL, Taiwan; USNM
366576, 1 female, 52 mm SL,01°12'S, 144°16’E, Ninigo

Atoll, Papua New Guinea; USNM 376173, 3 males and 3
females, 64-104 mm SL, 23°29’S, 152°05’E, One Tree
Island, Queensland, Australia; USNM 376171, 1 male and
1 female, 63 mm SL, 23°25’S, 151°55’E, Heron Island,
Queensland, Australia; USNM 366682, 1 female, 82
mm SL, 23°25’S, 151°55’E, Heron Island, Queensland,
Australia; USNM 366688, 1 male, 38 mm SL, Taiwan;
USNM 366693, 7 specimens, 21-82 mm SL, Taiwan.

Tentatively assigned specimens (A. aff. riukiuensis).
BPBM 40938, 5 males and 2 females,Taiwan; USNM
309643, 3 males and 3 females, 40-60 mm SL, 04°52’N,

119°26’E, Sibutu Island, Philippines.

Diagnosis. Vertebrae 11-12+30-32=41=44. dorsal fin
rays 77-92 (average 84), anal fin rays 59-72 (average 65);
eyes moderately large (2.5-3.0% SL); body moderately
slender, massive, snout with many cirri; scales on cheeks
and large scale patch above opercular spine (6-17), large
specimens with few scales also below opercular spine;
upper prcopercular pore present; outer pseudoclaspcr broad-
based, not very large, not extending beyond hood in resting
position; inner pseudoclaspcr not much smaller than outer
pseudoclasper, anterior and posterior lobes of equal size,
anterior lobe broadly connected to anterior rim of outer
pseudoclasper; otolith slender, length to height 2.2-2.4,
with gently curved dorsal rim, otolith length to sulcus length
1.6-1.7, ostium length to cauda length 3.5^1.0.

Description (Figs 17-19).The principal meristic and
morphometric characters are shown in Table 8. One of the
largest species attaining up to 150 mm SL; mature at about
50-55 mm SL. Body moderately slender, massive, with
moderately pointed head profile. Many cirri on snout. Eye
size 2.5-3.0 (3.0)% SL. Head with scale patch on cheek
containing 8-13 vertical rows of scales on upper part and
4-5 vertical rows on lower part, scale patch above opercular
spine with 6-17 scales, in adults, scaled patch below
opercular spine with up to 8 scales. Horizontal diameter
of scales on body of a 74 mm SL male, about 1.5% SL, in
about 40 horizontal rows. Maxillary ending far behind eye,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded with small knob. Anterior
nostril positioned high, 1/2-2.5 distance from upper lip to
anterior margin of eye. Posterior nostril small, about 1/6 to
1/8 the size of eye. Opercular spine with free tip, pointed.
Anterior gill arch with 15-21 rakers, 3 elongate. Small
plate-like raker between lower two elongate rakers in most
specimens. Pseudobranchial filaments 0-3 (usually 2).

Head sensory pores (Fig. 18A-D). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior): first anterior
pore large, tubular, with cirri. Preopercular pores: 3 lower,
first and second with joined opening, covered by dermal
flap in lateral view; third tubular; upper preopercular pore
present. [See description of Alionematichthys ceylonensis
for position of pores.]

115

W. Schwarzhans and P. R. Moller

Fig. 19. Alionematichthys a(T. riukiuensis (Aoyagi, 1954), A, lateral view ol head. BPBM 40938, male, 62 mm SL; B, ventral view ot head,
BPBM 40938, male, 62 mm SL; C, lateral view of head, USNM 309643, male, 58 mm SL; 1). inclined lateral view of male copulatory organ,
BPBM 40938, male, 80 mm SL; E, ventral view of male copulatory organ, BPBM 40938, male, 80 mm SL; F, view of left pseudoclaspers from
inside, USNM 309643,58 mm SL; G, view of left pseudoclaspers from inside, BPBM 40938, male, 62 mm SL; H, view ofleft pseudoclaspers
from inside, BPBM 40938, male, 80 mm SL; 1, median view of right otolith, USNM 309643, male, 58 mm SL.

116

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Dentition (of a 74 mm SL male, NSMT-P 55637).
Premaxilla with 6 outer rows of granular teeth and 2 inner
rows of larger teeth anteriorly. Anteriormost teeth in inner
row up to 1/2 diameter of pupil. Vomer horseshoe-shaped,
with 6 outer rows of small teeth and 1 inner row of large
teeth up to 1/4 diameter of pupil. Palatine with 4 outer
rows of small teeth and 1 inner row of long teeth up to 1/4
diameter of pupil. Dentary with 6 outer rows of granular
teeth and 1 inner row of larger teeth anteriorly, up to about
1/2 diameter of pupil.

Otolith (Fig. 18J). Elongate in shape, length to height
2.2-2.4 and thin (otolith height to otolith thickness about 3).
Anterior tip slightly pointed, posterior tip pointed, expanded
and irregularly ornamented. Dorsal rim gently curved,
without distinct angles. Inner face moderately convex, outer
face slightly concave to almost flat, both smooth. Otolith
length to sulcus length 1.6-1.7. Sulcus medianly positioned,
not inclined, with separated colliculi and marked notch at
ventral rim of sulcus at joint of ostium with cauda. Ostium
length to cauda length 3.5-4.0. Ventral furrow distinct and
close to ventral rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined and
5-8 depressed. Parapophyses present from vertebrae 6 to
11(12). Pleural ribs on vertebrae 2 to 11 (12). First anal fin
pterygiophore not or only slightly longer than subsequent
pterygiophore.

Male copulatory organ (Fig. 18E-I). Two pairs of
rather small pseudoclaspers, not extending beyond hood
in resting position. Outer pseudoclasper a broad based
flap; inner pseudoclasper slightly shorter, very thick (best
seen in ventral view: Fig. 18G), with anterior pointed lobe
broadly connected by ligament to anterior rim of outer
pseudoclasper, posterior lobe broad, about equal in size.
Isthmus moderately narrow; penis curved, slightly longer
than outer pseudoclaspers.

Colour. Live colour brownish-red or orange-brown
(SMNS 22804), sometimes with yellowish vertical fins
(Fig. 17A). Preserved colour medium to dark brown.

Variability. Alioneniatichthys riukiuensis shows some
variation in the number of scales on the opercle above and
below the opercular spine and a certain variation of the
dorsal and anal fin ray counts, both of which does not seem
to follow a geographic pattern.

Remarks. Machida (1994) described Dinematichthys
megasoma from Australia. These specimens agree well
with the variability observed in A. riukiuensis, including
the characters regarded by Machida as distinctive from
Dinematichthys riukiuensis, i.e. the short tubular posterior
nostril, the widely scaled opercle and the lateral scale row
counts. We have, however, observed two specific lots which
somewhat depart from the general range of variability
(see Fig. 19). They are both notable for few or no cirri on
the snout and few scales above the opercular spine (2-6)
(Fig. 19A-C). They vary amongst themselves in meristics
(see Table 1, 8). Their pseudoclaspers resemble those of
A. riukiuensis (Fig. 19D-H) perfectly as do their otoliths

(Fig. 191). Due to the limited amount of specimens available
and the high degree of similarity with A. riukiuensis they are
here referred to as Alioneniatichthys aff. riukiuensis.

Comparison. Alioneniatichthys riukiuensis belongs to
the species group with an upper preopercular pore, large
eyes and (many) scales above the opercular spine and cirri
on the snout (comprising also A. winterbottomi sp. nov.). It
differs mainly in the characters of the pseudoclaspers and
the presence of scales below the opercular spine in adults
(vs always absent in A. winterbottomi sp. nov.). It differs
from A. piger and A. samoaensis sp. nov., which likewise
show many cirri on the snout in the presence of many scales
above the opercular spine (vs none or rarely 1-2) and the
pseudoclasper morphology. Also, A. riukiuensis is notable
for the large size it can attain and at which it matures, greater
than in all other species of Alioneniatichthys.

Distribution. Alioneniatichthys riukiuensis is widely
distributed in the Indo-west Pacific from the Ryukyu
Islands, Hainan Island and the west coast ofThailand in the
north to the northern shores of Australia, New Caledonia
and Vanuatu in the south (Fig. 16).

Alioneniatichthys samoaensis sp. nov.

(Figs 16, 20, 21, Tables 1,8)

Material examined. (36 specimens, 22-59 mm SL).
HOLOTYPE - CAS 81486, male, 46 mm SL, ca. 11 °03'S,
171°04'W, seaward face of leeward reef on Swains Island,
American Samoa, coll. L. R. Taylor Jr., 23 May 1967.
PARATYPES-AMS IB. 6513, male, 54 mm SL, 14°27'S,
178°05'W,Alofi, Niue, W.N.McDowall, May 1963; BPBM
24122, 2 females, 56 and 59 mm SL, Samoa, Tutuila, R.
C. Wass, 1976-1977; CAS 227305,20 females, 22-53 mm
SL, 11 males, 25-47 mm SL, juvenile, 22 mm SL, same
data as holotype.

Diagnosis. Vertebrae 11-12+29-31=40-43, dorsal fin
rays 73-80, anal fin rays 57-64; eyes moderately large
(2.2-2.8% SL); body slender, snout with many cirri;
scales on cheeks, no scales above opercular spine; upper
preopercular pore present; posterior nostril funnel-shaped;
outer pseudoclasper broad-based, not extending beyond
hood in resting position, with thick distal knob on inner face;
inner pseudoclasper about half size of outer pseudoclasper.
anterior thorn slightly bent, posterior lobe broader than
anterior thorn.

Description (Figs 20, 21).The principal meristic and
morphometric characters are shown in Table 9. Body
moderately slender, with moderately pointed head profile.
Many cirri on snout. Eye size 2.2-2.8 (2.2)% SL. Head
with scale patch on cheeks containing 6-7 vertical rows of
scales on upper part and 2-3 vertical rows on lower part,
no scale patch on opercle. Horizontal diameter of scales
on body in holotype 1.7% SL, in about 24 horizontal rows.
Maxillary ending far behind eye, dorsal margin of maxillary
covered by upper lip dermal lobe, posterior end expanded.
Anterior nostril positioned high, one-half distance from
upper lip to anterior margin of eye. Posterior nostril with

117

W. Schwarzhans and P. R. Moller

Fig. 20. Alionematichthys samoaensis sp. nov., CAS 81486, holotype, male, 46 mm SL.

Fig. 21. Alionematichthys samoaensis sp. nov., A. lateral view of head, holotype; B, ventral view of head, holotype; C, inclined lateral view
of male copulatory organ. CAS 81486. male, 39 mm SL; D, ventral view of male copulatory organ, holotype; E, view of left pscudoclaspers
from inside, holotype; F, view ofleft pseudoclaspers from inside, CAS 81486, 39 mm SL.

broad funnel-shaped rim, about one-eighth the size of eye.
Opercular spine with free tip, pointed. Anterior gill arch
with 14-18 (15) rakers, 3 elongate. In holotype and a few
paratypes lower two elongate rakers interrupted by one
plate-like raker. In other paratypes, three elongate rakers in
row. Pseudobranchial filaments 1-2 (usually 2).

Head sensory pores (Fig. 21 A, B). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third the size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior); first
anterior pore large, tubular, with cirri. Preopercular pores:
3 lower, first and second with joined opening, covered by

118

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Table 9. Meristic and morphometric characters of Alionematichthys
samoaensis sp. nov.

Holotype

CAS

81486

Holotype +

35 non-types

Mean (range)

Standard length in mm

37.3 (22-59)

Meristic characters

Dorsal fin rays

76.6 (73-80)

Caudal fin rays

16.1 (16-17)

Anal fin rays

59.4 (57-64)

Pectoral fin rays

21.6(20-22)

Precaudal vertebrae

11.2(11-12)

Caudal vertebrae

30.7(29-31)

Total vertebrae

41.9(40-43)

Rakers on anterior gill arch

15.6(14-18)

Pseudobranchial filaments

1.9 (1-2)

D/V

5.6 (5-6)

D/A

22.6 (20-26)

V/A

13.2(13-14)

Morphometric characters in % of SL

Head length

27.2

27.2 (26.2-28.9)

Head width

14.2

13.8(11.8-16.2)

Head height

16.4

16.6(15.3-17.9)

Snout length

5.5

6.1 (5.5-6.5)

Upper jaw length

14.7

14.3(13.3-15.6)

Diameter of pigmented eye

2.2

2.5 (2.2-2.8)

Diameter of pupil

1.4

1.4 (1.2-1.5)

Interorbital width

7.8

7.6 (7.2-8.0)

Posterior maxilla height

4.7

4.7 (3.9-5.3)

Postorbilal length

19.8

19.4(18.5-21.2)

Preanal length

48.7

47.7 (44.8-50.4)

Predorsal length

30.7

31.1 (28.9-32.5)

Body depth at origin of anal fin

19.4

18.8(16.3-20.4)

Pectoral fin length

13.4

13.5(12.7-15.7)

Pectoral fin base height

6.5

6.6 (6.2-6.8)

Ventral fin length

26.0

25.5 (22.5-27.1)

Base ventral fin - anal fin origin

29.6

28.3(25.8-31.6)

dermal flap in lateral view; third tubular; upper preopercular
pore present. [See description of Alionematichthys
ceylonensis for position of pores.]

Dentition (of holotype). Premaxilla with 5 outer
rows of granular teeth and 1 inner row of larger teeth
anteriorly. Anteriormost teeth in inner row up to 1/3
diameter of pupil. Vomer horseshoe-shaped, with 2 outer
rows of small teeth and 1 inner row of large teeth up to
1/3 diameter of pupil. Palatine with 1 outer row of small
teeth and 1 inner row of long teeth up to 1/3 diameter of
pupil. Dentary with 5 outer rows of granular teeth and
1 inner row of larger teeth anteriorly, up to about 2/3
diameter of pupil.

Otolith (not known).

Axial skeleton. Neural spine of vertebra 4 inclined and
5-8 depressed. Parapophyscs present from vertebrae 6 to 11
(12). Pleural ribs on vertebrae 2 to 10-11 (12). First anal fin
pterygiophore not or only slightly longer than subsequent
pterygiophore in females, much longer in males, but not
reaching last precaudal parapophysis.

Male copulatory organ (Fig. 21C-F). Two pairs of rather
small pseudoclaspers, not extending beyond hood in resting
position. Outer pseudoclasper a broad based flap, with
thick knob distally on inner face, its supporter somewhat
bent backward at tip; inner pseudoclasper thick, about half
the size of outer pseudoclasper, anterior lobe slightly bent,
posterior lobe broader than anterior thorn. Isthmus narrow;
penis curved, slightly longer than outer pseudoclaspers.

Colour. Live colour unknown. Preserved colour
uniformly light brown.

Comparison. Alionematichthys samoaensis belongs to
the species group with an upper preopercular pore, large
eyes and no scales above the opercular spine, comprising
also A. piger and A. suluensis sp. nov. Alionematichthys
samoaensis obviously is closely related to A. piger, which
it replaces in region of Samoa, differing mainly in the
distinct knob on the inner face of the outer pseudoclasper
(vs thin outer pseudoclasper without knob) and the funnel-
shaped posterior nostril (vs anterior rim elevated only).
From A. suluensis, it differs readily in the presence of many
delicate cirri on the snout (vs absent) and the anterior lobe
of the inner pseudocasper being as long as its posterior lobe
(vs twice as long).

Fig. 22. Alionematichthys shinoharai sp. nov., NSMT-P 34895, holotype, male, 34 mm SL.

119

W. Schwarzhans and P. R. Moller

Fig. 23. Alionematichthys shinoharai sp. nov., A, lateral view ofhcad, holotype; B, ventral view of head, holotype; C, view of left pseudoclaspers
from inside, holotype; D, inclined lateral view of male copulatory organ, holotype; E, median view of right otolith, holotype; F, ventral view
of right otolith, holotype; G, median view of right otolith, YCM-P36416, female, 42 mm SL.

Distribution. Alionematichthyssamoaensis is endemic
to the Islands of the Samoa group, and is in fact the
only endemic Dinematichthyini restricted to that region
(Fig. 16).

Biology. A 46 mm SL female (CAS 81486) contains ca.
100 embryos, length 4.6 mm TL,

Each embryo with three short pigmented rows on the
posterior part of the body.

Etymology. Named after the type locality, American
Samoa. It is an adjective.

Alionematichthys shinoharai sp. nov.

(Figs 1,22,23, Tables 1, 10)

Material examined. (2 specimens, 34^12 mm SL).
HOLOTYPE - NSMT-P 34895, male, 34 mm SL,
28°H’2”N, 129°16’E, Sakinome beach, Amami-Oshima
Island, Ryukyu Islands, Japan, 5 m, K. Matsuura and M.
Aizawa, 14 June 1991. PARATYPES - YCM-P 36416,
female, 42 mm SL, Amami Islands, Kakeroma Island,
Ryukyu Islands, Japan, 24 Aug. 1995.

Diagnosis. Vertebrae 11+32-33=43-44, dorsal fin rays
83-85, anal fin rays 67; eyes large (3.0-3.3% SL); body
slender, snout pointed, without cirri; scales on cheeks,
two scales above opercular spine; upper preopcrcular
pore absent; outer pseudoclasper broad-based simple flap;
inner pseudoclasper very small, somewhat depressed;
otolith moderately elongate, length to height 2.1 -2.2, with
regularly curved dorsal rim, sulcus small, otolith length
to sulcus length 1.9-2.0, cauda very small, ostium length
to cauda length 5.5-7, ostium height to cauda height
2 . 2 - 2 . 7 .

Description (Figs 22-23).The principal meristic and
morphometric characters are shown in Table 10. Mature at

about 35-40 mm SL (male of 34 mm SL probably subadult).
Body slender, with pointed head profile. No cirri on snout.
Eye size 3.3 (3.0)% SL. Head with scale patch on cheeks
containing 7 vertical rows of scales on upper part and 4
vertical rows on lower part, 2 scales on opercle above
opercular spine. Horizontal diameter of scales on body of
holotype 1.5% SL, in about 24 horizontal rows. Maxillary
ending far behind eye, dorsal margin of maxillary covered
by upper lip dermal lobe, posterior end expanded with small
knob. Anterior nostril positioned high, one-third distance
from upper lip to anterior margin of eye. Posterior nostril
with narrow elevated rim, about one-eighth the size of eye.
Opercular spine with free tip, sharply pointed. Anterior
gill arch with 15(16) rakers, 3 elongate. Pseudobranchial
filaments 2.

Head sensory pores (Fig. 23 A, B). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third the size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior): first
anterior pore large, tubular, with single cirrus anteriorly.
Preopercular pores: 3 lower, first and second with joined
opening, covered by dermal flap in lateral view; third
tubular; upper preopercular pore absent. [See description
of Alionematichthys ceyionensis for position of pores.]

Dentition (of holotype). Premaxilla with 5 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 1 /4 diameter of pupil.
Vomer horseshoe-shaped, with 1 outer row of small teeth
and 1 inner row of larger teeth up to 1/5 diameter of pupil.
Palatine with 1 outer row of small teeth and 1 inner row
of longer teeth up to 1/4 diameter of pupil. Dentary with 5

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Dinematichthyine fishes of the Indo-west Pacific, Part IV

Table 10. Meristic and morphometric characters of Alionematichthys
shinoharai sp. nov.

Holotype

NSMT-

P34895

Paratype

YCM-

P36416

Standard length in mm

Meristic characters

Dorsal fin rays

Caudal fin rays

Anal fin rays

Pectoral fin rays

Precaudal vertebrae

Caudal vertebrae

Total vertebrae

Rakers on anterior gill arch

Pseudobranchial filaments

D/V

D/A

VIA

Morphometric characters in % of SL

Head length

27.2

26.7

Head width

11.9

12.8

Head height

15.9

16.2

Snout length

5.6

6.5

Upper jaw length

13.8

Diameter of pigmented eye

3.0

3.3

Diameter of pupil

2.0

2.1

Interorbital width

6.5

7.0

Posterior maxilla height

4.0

Postorbital length

19.8

18.0

Preanal length

44.8

44.9

Predorsal length

32.0

32.6

Body depth at origin of anal fin

16.1

18.1

Pectoral fin length

14.1

15.1

Pectoral fin base height

6.6

5.5

Ventral fin length

24.7

22.6

Base ventral fin - anal fin origin

25.2

24.1

outer rows of granular teeth and 1 inner row of larger teeth
anteriorly, up to about 1/3 diameter of pupil.

Otolith (Fig. 23E-G). Moderately elongate in shape,
length to height 2.1-2.2 (34-40 mm SL), rather thick
(otolith height to otolith thickness about 2). Anterior tip
sharply pointed, posterior tip pointed, slightly expanded.

Dorsal rim gently curved, with postdorsal angle in male
specimen (Fig. 23E) and absent in female (Fig. 23G),
exhibiting slight sexual dimorphism. Inner and outer face
moderately convex, smooth. Otolith length to sulcus length
1.9-2.0. Sulcus medianly positioned, slightly inclined, with
separated colliculi, without marked notch at ventral rim
of sulcus at joint of ostium with cauda. Cauda very small,
ostium length to cauda length 5.5-7, ostium height to cauda
height 2.2-2.7. Ventral furrow feeble, close to ventral rim
of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined and
5-8 depressed. Parapophyses present from vertebrae 6 to 11.
Pleural ribs on vertebrae 2 to 10. First anal fin pterygiophore
only slightly longer than subsequent pterygiophore.

Male copulatory organ (Fig. 23C, D). Two pairs of
small pseudoclaspers, which may not be fully mature in
single male specimen investigated. Outer pseudoclasper a
simple flap with broad base; inner pseudoclasper a small
depressed flap without significant indentation. Isthmus
moderately narrow; penis slightly curved, longer than outer
pseudoclaspers.

Colour. Live colour unknown. Preserved colour
uniformly light brown.

Comparison. Alionematichthys shinoharai belongs
to the species group without an upper preopercular
pore together with A. ceylonensis, A. phuketensis and
A. plicatosurculus. It differs from all these in the presence
of two scales above the opercular spine (vs none), the
somewhat less highly positioned anterior nostril with one-
third distance from upper lip to anterior margin of eye (vs
1/2-2.5) and the peculiar otolith with its short sulcus and the
very small cauda. The combination of these characters is so
distinctive that they could indicate a separate genus, which,
however, will be subject to review of a larger male specimen
in order to fully evaluate the pseudoclasper pattern.

Distribution. Alionematichthys shinoharai has so far
only been found at the northern islands of the Ryukyu Island
chain of Japan, i.e. Amami-Oshima (Fig. I).

Etymology. Named in honour of Gento Shinohara,
NSMT, Tokyo, in recognition of his many contributions to
ichthyology and for his kind support of the present revision.
It is a noun in apposition.

Fig. 24. Alionematichthys suluensis sp. nov., AMS 1.40149-028, holotype, male, 40 mm SL.

121

W. Schwarzhans and P. R. Moller

Alionematichthys suluensis sp. nov.

(Figs 9,24, 25, Tables 1, 11)

Material examined. (57 specimens, 26-49 mm SL).
HOLOTYPE - AMS 1.40149-028, male, 40 mm SL,
12°21.70N, 121°27.57E, 0-20 m, rocky surge area at base
of cliffs, S tip of Buyamao Island off SE Mindoro Island,
Philippines, coll. MIN team, rotenone, 30 May 2000.
PARATYPES - AMS 1.40149-028), 1 male, 30 mm SL, 5
females, 30-42 mm SL, 3 juveniles, 13-17 mm SL, same
data as holotype; USNM 263691, 1 male, 43 mm SL and
5 females, 41-45 mm SL, 08°5r42”N, 123°24’36”E,
Solino Island, Zamboanga, Mindanao, Philippines, 0-5 m,
A. Alcala et al., 4 May 1979; USNM 263694, 6 males and
12 females, 27-49 mm SL, 10°52’N, 121°12’E, Palawan,
Philippines; USNM 376216,4 males and 1 female, 29-45
mm SL, 09°04'N, 123°08'E, Negros, Philippines; USNM
366601,1 male, 46 mm SL, 09°04’N, 123°16’E,Apo Island,
Philippines; USNM 366602,1 female, 34 mm SL, 09°23’N,
123°15’E, Negros, Philippines; USNM 374183, 1 male,
31 mm SL, 10°35’05”N, 122°08’30”E, rocky tidepool,
Talisayan Point, San Joaquin, Lawigan, Iloilo Province,

Philippines, 0-7 m, J. T. Williams et al., 25 Sep. 1995;
USNM 376179, 3 males and 6 females, 26-48 mm SL,
09°03’N, 122°59’E, Negros, Philippines; USNM 376181 ’
3 males, 37-44 mm SL, 10°55’05”N, 121°02’03”E, Putic
Island, Palawan, Philippines, 0-4.6 m, V. Springer et al. t
22 May 1978; USNM 376182,2 males, 34 and 44 mm SL,’
09°06’30”N, 122°55’24”E, near Giligaon, north of Maloh,
Negros. Philippines, 0-2 m, L. W. Knapp et al., 26 April
1979; USNM 376186, 2 males, 32 and 34 mm SL and ]
female, 41 mm SL, 08°51’24”N, 123°24’36”E, Solino
Island, Zamboanga, Mindanao, Philippines, 0-5 m, L. W.
Knapp et al., 3 May 1979.

Diagnosis. Vertebrae 1 1-12+30-32=41-43, dorsal fin
rays 72-82, anal fin rays 56-64; eyes moderately large
(2.1—2.5% SL); body slender, snout without cirri; scales
only on cheeks, no scales above opercular spine; upper
preopercularpore present; outer pseudoclasper moderately
broadbascd, long, sometimes extending beyond hood in
resting position; inner pseudoclasper half as long as outer
pseudoclasper, anterior lobe very long, twice as long as
posterior lobe; otolith length to height 2.1-2.2, with shallow

Fig. 25. Alionematichthys suluensis sp. nov.. A, lateral view of head, holotype; B, ventral view of head, holotype; C, view of left pseudoclaspers
from inside, holotype; D, inclined lateral view of male copulatory organ, holotype; E, median view of right otolith, AMS 1.40149-028, female,
40 mm SL; F, ventral view of right otolith, AMS 1.40149-028, female, 40 mm SL.

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Dinematichthyine fishes of the Indo-west Pacific, Part IV

Table 11. Meristic and morphometric characters of Alionematichthys
suluensis sp. nov.

Holotype
AMS I.
40149-
028

Holotype +

56 paratypes

Mean (range)

Standard length in mm

36.7(13-48)

Meristic characters

Dorsal fin rays

77.5 (72-82)

Caudal fin rays

15.8(15-16)

Anal fin rays

60.2 (56-64)

Pectoral fin rays

20.2(19-22)

Precaudal vertebrae

11.5(11-12)

Caudal vertebrae

30.7 (30-32)

Total vertebrae

42.1 (41-43)

Rakers on anterior gill arch

14.5(12-16)

Pseudobranchial filaments

D/V

6.1 (6-7)

D/A

22.3 (20-25)

V/A

13.6(13-15)

Morphometric characters in % of

Head length

24.4

25.2 (23.9-27.0)

Head width

12.2

11.9(10.2-13.6)

Head height

14.5

15.1 (14.3-16.6)

Snout length

5.7

5.5 (4.9-6.1)

Upper jaw length

12.3

13.0(12.3-13.9)

Diameter of pigmented eye

2.2

2.2 (2.1-2.5)

Diameter of pupil

1.2

14(1.2-1.7)

Interorbital width

6.3

6.6 (6.0-7.4)

Posterior maxilla height

4.0

4.2 (3.7-4.7)

Postorbilal length

16.9

18.0 (16.9-18.9)

Preanal length

46.0

46.5 (43.6-48.6)

Predorsal length

31.0(29.4-32.5)

Body depth at origin of anal fin

15.7

15.8(13.5-18.4)

Pectoral fin length

12.1

12.9(10.8-14.5)

Pectoral fin base height

4.4

5.3 (44-5.8)

Ventral fin length

25.6

24.0(21.2-25.9)

Base ventral fin - anal fin origin

28.6

28.4(26.0-31.6)

dorsal rim and postdorsal angle, otolith length to sulcus
length 1.7-1.8, ostium length to cauda length 3.3-3.7.

Description (Figs 24, 25).Thc principal meristic and
morphometric characters are shown in Table 11. One of
the smallest species in the genus; mature at about 30 mm
SL, maximum size about 50 mm SL. Body slender, with
moderately pointed head profile. No cirri on snout. Eye
size 2.1-2.5 (2.2)% SL. Head with scale patch on cheek
containing 5-7 (5) vertical rows of scales on upper part
and 3 vertical rows on lower part, no scales on opercle.
Horizontal diameter of scales on body about 2.0% SL,
in 24 horizontal rows. Maxillary ending far behind eye,
dorsal margin of maxillary covered by upper lip dermal
lobe, posterior end expanded with little knob. Anterior
nostril positioned high, one-third distance from upper lip to
anterior margin of eye. Posterior nostril small, about one-

sixth the size of eye. Opercular spine with free tip, pointed.
Anterior gill arch with 12-16 (14) rakers, 3 elongate in a
row. Pseudobranchial filaments 2.

Head sensory ’ pores (Fig. 25A, B). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about half the size of three anterior pores.
Mandibular pores 6 (3 anterior and 3 posterior): first
anterior pore large, tubular, with single cirrus anteriorly.
Preopercular pores: 3 lower, first and second with joined
opening, covered by dermal flap in lateral view; third
tubular; upper preopercular pore present. [See description
of Alionematichthys ceylonensis for position of pores.]

Dentition (of holotypc). Premaxilla with 4 outer rows
of granular teeth and 1 inner row of larger teeth anteriorly.
Anteriormost teeth in inner row up to 3/4 diameter of pupil.
Vomer horseshoe-shaped, with 1 outer row of small teeth
and 1 inner row of larger teeth up to 1/3 diameter of pupil.
Palatine with an outer row of small teeth and an inner row
of larger teeth up to 1/3 diameter of pupil. Dentary with 3
outer rows of granular teeth and 1 inner row of larger teeth
anteriorly, merging into 1 row of larger teeth posteriorly.
Large dentary teeth up to about 3/4 diameter of pupil.

Otolith (Fig. 25E, F). Moderately elongate in shape,
length to height 2.1-2.2 (30-49 mm SL) and moderately
thick (otolith height to otolith thickness about 1.5). Anterior
tip slightly pointed, posterior tip pointed, expanded and
irregularly ornamented. Dorsal rim shallow, with notch
above anterior tip and behind marked postdorsal angle.
Inner face convex, outer face slightly concave to almost flat,
both smooth. Otolith length to sulcus length 1.7-1.8. Sulcus
slightly supramedially positioned and slightly inclined,
with separated colliculi and marked notch at ventral rim
of sulcus at joint of ostium with cauda. Ostium length to
cauda length 3.3—3.7. Ventral furrow very close to ventral
rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined
and 5-8 depressed. Parapophyses present from vertebrae
6 to 11 (12). Pleural ribs on vertebrae 2 to 11 (12). First
anal fin pterygiophore markedly longer than subsequent
pterygiophore, reaching tip of last precaudal parapophysis
in males.

Male copulatory organ (Fig. 25C, D). Two pairs of
pseudoclaspers, outer sometimes extending beyond hood
in resting position. Outer pseudoclasper moderately broad
based with thick supporter bent backwards at tip; inner
pseudoclasper about half the length, thin, with very long
anterior inclined lobe about twice as long as small postci ioi
lobe. Isthmus narrow; penis curved, slightly shorter than
outer pseudoclaspers.

Colour. Live colour unknown. Preserved colour
uniformly brown.

Comparison. Alionematichthys suluensis belongs to
the species group with an upper preopercular pore, large
eyes and no scales above the opercular spine as A. piger
and A. samoaensis. It differs from both in the lack of cini
on the snout (except for a single cirrus associated with the

123

W. Schwarzhans and P. R. Moller

Fig. 26. Alionematichthys winterbottomi sp. nov., ROM 47597, holotype, male, 54 mm SL.

Fig. 27. Alionematichthys winterbottomi sp. nov.. A, lateral view of head, ROM 47597, holotype, male, 54 mm SL; B, ventral view of head,
holotype; C, ventral view of male copulatory organ, ROM 82975, paratype, 58 mm SL; I), inclined lateral view of male copulatory organ, ROM
82975, paratype, 58 mm SL; E, view of left pseudoclaspers from inside, ROM 82975, paratype, 65 mm SL; F, view of left pseudoclaspers
from inside, holotype; G, median view of right otolith, CAS 227284, male, 76 mm SL.

124

Dinematichthyine fishes of the Indo-west Pacific, Part IV

first anterior mandibular pore) and the very peculiar shape
of the inner pseudoclasper with its long anterior thorn.

Distribution. Alionematichthys suluensis appears to
be restricted in distribution to the Philippines between 13°
and 8° N, mainly in the Sulu Sea, where it co-occurs with
A. piger, A. plicatosurculus and A. riukiuensis (Fig. 16).

Biology. A 39 mm SL female (USNM 376179), contains
325 embryos, length 3.7 mm TL.

Etymology. Named after the type locality, the Sulu Sea
of the Philippines. It is an adjective.

Alionematichthys winterhottomi sp. nov.

(Figs 16, 26, 27, Tables 1, 12)

Material examined. (217 specimens, 10-88 mm SL).
HOLOTYPE - ROM 47597, male, 80 mm SL, 18°45 , 52”S,

178°31’ 13 ’’ E, shallow reef off first black rock 300 m S of
University of the South Pacific Research Station, Dravuni
Island, Fiji, coll. R. Winterbottom, A. R. Emery, F. Emery
and R. McKinnon, 20 March 1983. PARATYPES - CAS
222530, 2 males, 56 and 57 mm SL and 1 female, 42 mm
SL, Vatumatau Bay, Fiji, D. W. Greenfield et al., 12 Nov.
2002; CAS 227291, 1 male, 65 mm SL and 1 female, 57
mm SL, Viti Levu north, Fiji, D. W. Greenfield et al., 31
March 2002; CAS 222532, 1 male, 48 mm SL, 1 female,
49 mm SL and 1 juvenile, 29 mm SL, Fiji, D. W. Greenfield
et al., 14 Nov. 2002; CAS 222540, 1 female, 52 mm SL,
Fiji, 3 April 2002; CAS 222544, 2 males, 42 and 48 mm
SL and 2 females, 45 and 55 mm SL, Fiji, D. W. Greenfield
et al., 14 March 1982; CAS 222552, 4 males, 51-70 mm
SL, 2 females, 48-54 mm SL, and 2 juveniles, 22-23 mm
SL, Great Sea reef, Kia Island, Fiji, D. W. Greenfield et al.
31 March 2002; CAS 222558, 1 male, 63 mm SL, and 1
juvenile, 35 mm SL, Fiji, D. W. Greenfield et al. 15 Nov.
2002; CAS 227284,7 males, 54-75 mm SL and 5 females,
41-64, Fiji, reef NE ofYaqaga Island and NW of Ovatova
reef, D. W. Greenfield et al. 25 March 2002; CAS 227307,
3 males, 42-57 mm SL, 3 females, 40-59 mm SL, Bua Bay,
Fiji, D. W. Greenfield et al. 24 March 2002; CAS 222586,
2 males, 42-44 mm SL and 3 females, 43, 50 and 50 mm
SL, Nasau Bay, Vanua Levu, Fiji, D. W. Greenfield et al.,
25 May 2003; CAS 222589, 1 female, 57 mm SL, Fiji, D.
W. Greenfield et al. 25 May 2003; CAS 222590, 2 males,
51 and 65 mm SL, 1 juvenile, 25 mm SL; Fiji east shore, D.
W. Greenfield et al. 15 March 2002; CAS 222592, 1 male,
46 mm SL, 2 females, 31 and 44 mm SL, Budd Reef, Fiji,
D. W. Greenfield et al. 23 May 2003; CAS 227294, 1 male,
57 mm SL, Fiji, D. W. Greenfield et al., 20 May 2003; CAS
227295, 4 males, 45-57 mm SL, 12 females, 29-58 mm
SL, Budd Reef, Fiji, D. W. Greenfield et al., 22 May 2003;
CAS 227297, 1 male, 53 mm SL and 3 females, 35-52
mm SL, Fiji, D. W. Greenfield et al, 23 May 2003; CAS
222605,2 females, 44 and 73 mm SL, Fiji, D. W. Greenfield
et al., 5 January 2003; CAS 227298, 1 female, 88 mm SL,
Fiji, D. W. Greenfield et al, 28 May 1999; CAS 227299, 3
males, 55-75 mm SL and 1 female, 53 mm SL, Fiji, D. W.
Greenfield et al, 29 January 2002; CAS 222608, 3 males,

Table 12. Meristic and morphometric characters of Alionematichthys
winterhottomi sp. nov.

Holotype

ROM

47597

Holotype +

198 paratypes

Mean (range)

Standard length in mm

48.9(10-88)

199

Meristic characters

Dorsal fin rays

82.5 (80-86)

Caudal fin rays

Anal fin rays

63.9 (61-65)

Pectoral fin rays

21.3 (20-22)

Precaudal vertebrae

11.0(11-12)

Caudal vertebrae

31.0(29-32)

Total vertebrae

42.0 (40-43)

Rakers on anterior gill arch

18.3(17-20)

Pseudobranchial filaments

1.9 (1-2)

D/V

5.7 (5-6)

D/A

23.1 (21-24)

VIA

Morphometric characters in % of SL

Head length

27.2

27.3 (26.3-27.8)

Head width

15.9

15.5(14.0-17.2)

I lead height

16.4

16.9(15.9-18.3)

Snout length

5.2

6.0 (5.2-6.5)

Upper jaw length

14.4

14.4(13.6-15.1)

Diameter of pigmented eye

2.2

2.5 (2.2-3.0)

Diameter of pupil

1.4

1.5 (1.2-1.7)

Intcrorbital width

6.8

7.1 (6.2-7.9)

Posterior maxilla height

5.0

4.9 (4.5-5.4)

Postorbital length

20.4

20.0(19.5-20.6)

Prcanal length

47.1

46.9 (44.7-48.6)

Predorsal length

30.7

31.5(30.4-32.5)

Body depth at origin of anal fin

19.7

18.2(17.2-19.7)

Pectoral fin length

15.3

14.6(13.2-15.5)

Pectoral fin base height

6.7

6.6 (6.3-7.2)

Ventral fin length

22.8

23.0(21.4-24.4)

Base ventral fin - anal fin origin

26.3

26.7 (23.4-29.5)

38-44 mm SL, Fiji, D. W. Greenfield et al., 7 February
2002; CAS 222613, 3 males, 43-60 mm SL, Barrier reef
of Suva, Fiji, D. W. Greenfield et al. 27 Jan. 2002; CAS
222619, 1 male, 64 mm SL, Fiji, D. W. Greenfield et al,
31 May 1999; CAS 222621, male, 46 mm SL, female, 50
mm SL, Nukaiau Island, off Suva, D. W. Greenfield et al.,
28 Jan. 200?; ROM 82975, 26 males, 34-78 mm SL, 57
females, 22-75 mm SL, 16 juveniles, 10-34 mm SL, same
data as holotype; ROM 82977,5 females 45-70 mm SL, 2
males, 60 and 77 mm SL, 18°46’3(TS, I78°30’28”E, off
Yanu-Yanu-Sau Island, south of Dravuni, Fiji, A. R. Emery
and party, 30 Mar. 1983.

Additional material. USNM 377255, 15 specimens,
Vuro, Fiji; USNM 378376, 4 specimens, 27-50 mm SL,
Malolo, Fiji; USNM 257649, 1 male and 2 females,
32-58 mm SL, 17°ITS, 176°54’E, YasawaGroup, Viwa
Island, Fiji; USNM 263659, 2 males and 2 females,

125

W. Schwarzhans and P. R. Moller

30-72 mm SL, 18°52’S, 178°30’E, Kandavu Island, Fiji;
USNM 374211, 1 male, 65 mm SL, 21°20’S, 174°58’W,
Eua, Tonga; USNM 366566, 9 specimens, 36-77 mm
SL, 18°50’S, 178°32’E, Kandavu Island, Fiji; USNM
376196, 2 females, 39-48 mm SL, 18°56’S, 178°21’W,
Lau Group, Yagasa Island, Fiji; USNM 376209, 1 male
and 1 female, 45-49 mm SL, 17°06’S, 177°13’E, Yasawa
Group, Naviti Island, Fiji.

Diagnosis. Vertebrae ll(-12)+29-32=40-43, dorsal
fin rays 80-86, anal fin rays 61-65; eyes moderately large
(2.2-3.0% SL); body moderately slender, snout with many
cirri; scales on cheeks, patch of scales above opercular spine
with 5-9 scales; upper preopercular pore present; outer
pseudoclasper broad-based, not extending beyond hood in
resting position; inner pseudoclasper short, thin, free from
outer pseudoclasper, bifurcate, anterior pointed lobe short,
posterior lobe much expanded posteriorly; otolith length to
height 2.0-2.2, with shallow dorsal rim and weak postdorsal
angle, otolith length to sulcus length 1.6-1.7, ostium length
to cauda length 3.CM.5.

Description (Figs 26, 27).The principal meristic and
morphometric characters are shown in Table 12. Mature at
about 45 mm SL. Body moderately slender, with pointed
head profile. Many cirri on snout. Eye size 2.2-3.0 (2.2)%
SL. Head with scale patch on cheek containing 8-10 (9)
vertical rows of scales on upper part and 4 vertical rows
on lower part, 5-9 (9) scales on opercle above opercular
spine. Horizontal diameter of scales on body in holotypc
about 1.9% SL, in 30 horizontal rows. Maxillary ending far
behind eye, dorsal margin of maxillary covered by upper
lip dermal lobe, posterior end expanded. Anterior nostril
positioned high, 1/2.5 distance from upper lip to anterior
margin of eye. Posterior nostril small, about one-quarter the
size of eye. Opercular spine with free tip, pointed. Anterior
gill arch with 17-20 (17) rakers, Upper branch with one
knob-like raker and 3-5 (3) plate-shaped rakers and lower
branch with 13-15 (13) rakers, all plate-shaped except for
knob-shaped first and third. Pseudobranchial filaments 1-2
(usually 2).

Head sensory pores (Fig. 27A, B). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior): first
anterior pore large, tubular. Preopercular pores: 3 lower,
first and second with joined opening, covered by dermal
flap in lateral view; third tubular; upper preopercular pore
present. [See description of Alionematichthys ceylonensis
for position of pores.]

Dentition (of holotype). Premaxilla with 7 outer rows
of granular teeth and 2 inner rows of larger teeth anteriorly.
Anteriormost teeth in inner row up to 1/3 diameter of pupil.
Vomer horseshoe-shaped, with 3 outer rows of small teeth
and 1 inner row of large teeth up to 1/3 diameter of pupil.
Palatine with 3 outer rows of small teeth and 1 inner row
of 8 large teeth up to 1/3 diameter of pupil. Dentary with 5
outer rows of granular teeth and 1 inner row of larger teeth

anteriorly, merging into 1 row of larger teeth posteriorly.
Thirteen large dentary teeth, up to about 2/3 diameter of
pupil.

Otolith (Fig. 27G). Moderately elongate in shape,
length to height 2.0-2.2 and moderately thin (otolith
height to otolith thickness about 2.2-2.8). Anterior tip
slightly pointed, posterior tip pointed, expanded and
irregularly ornamented. Dorsal rim shallow, with notch
above anterior tip and with indistinct postdorsal angle.
Inner face convex, outer face slightly concave to flat, both
smooth. Otolith length to sulcus length 1.6-1.7. Sulcus
slightly supramedially positioned and slightly inclined,
with separated colliculi and marked notch at ventral rim
of sulcus at joint of ostium with cauda. Ostium length
to cauda length 3.0-4.0 (4.5). Ventral furrow weak, very
close to ventral rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined and
5-8 depressed. Parapophyses present from vertebrae 6 to
11 (12). Pleural ribs on vertebrae 2 to 11 (12). First anal fin
ptcrygiophorc slightly to markedly longer than subsequent
pterygiophore, but not reaching tip of last precaudal
parapophysis.

Male copulatory organ (Fig. 27C-F). Two pairs of
pseudoclaspers, outer not extending beyond hood in
resting position. Outer pseudoclasper broad based, nearly
triangular in shape, somewhat thickened towards lip; inner
pseudoclasper less half the length, thin, bifurcate, anterior
pointed lobe short, anteriorly free from outer pseudoclasper
halfway up from base, posterior lobe much expanded
posteriorly. Isthmus narrow; penis curved, slightly longer
than outer pseudoclaspers.

Colour. After preservation the specimens are uniformly
brownish.

Comparison. Alionematichthys winterbottomi belongs
to the species group with an upper preopercular pore, large
eyes, many cirri on the snout and within that group to the
subgroup with 5 or more scales above the opercular spine
together with A. riukiuensis, from which it differs in the
specific shape of the thin, free and small inner pseudoclasper
with its bifurcate shape and the lack of scales below the
opercular spine. Both species arc obviously is closely
related to each other.

Distribution. Alionematichthys winterbottomi is
restricted in distribution to the Fiji and Tonga Islands, where
it replaces A. riukiuensis (Fig. 16).

Etymology. Named in honour of Richard Winterbottom,
ROM, Toronto, Canada, in recognition of his many
contributions to ichthyology and for his great support of
the present revison. It is a noun in apposition.

Alionematichthys sp. 1
(Figs 16, 28, Table 1)

Material examined. (2 specimens, 75-87 mm SL).
AMS 19108-032, 1 male, 75 mm SL, 14°40’S, 145°28’E,
Lizard Island, Great Barrier Reef, Australia, 1-10 m, D.
F. Hoese and party, 17 Nov. 1975; WAM R30305-036, 1

126

Dinematichthyine fishes of the Indo-west Pacific, Part IV

female, 87 mmSL, 13°52’S, 126°56’E, Sir Graham Moore
Islands, Western Australia, depth 0-1 m, G. R. Allen, 15
Aug. 1991.

Remarks. Two specimens from northern Australia are
characterised by the presence of a uniform scale patch on
the opercle reaching from above the opercular spine to well
below it. The single male shows an inner pseudoclasper
resembling that of A. riukiuensis except for the anterior
rim of the inner pseudoclasper being free of a connection
to the outer pseudoclasper. All other characters, such as
cirri on the snout, head pores, otolith measures, meristics
and morphometries perfectly fit within the variation of
A. riukiuensis. More material needs to be studied before it
can be concluded if these specimens represent an extreme
variant of A. riukiuensis or a new species.

Alionematichthys sp. 2

(Figs 16, 29, Table 1)

Material examined. (1 specimen, 83 mm SL). WAM
P.25111-047, 1 female, 83 mm SL,20°28’30”S, 116°32’E,

Australia, Kendrew Island, Dampier Arch., G. R. Allen and
R. Steene, 3. Nov. 1974.

Remarks. A single female specimen from NW
Australia likely represents an undescribed species which
is characterised by the absence of cirri on the snout, the
absence of an upper preopercular pore (transformed to a
wart) and the presence of two scales above the opercular
spine. A similar combination of characters is found in
A. shinoharai, from which it differs in the more stout
and massive body and head and the longer sulcus on the
inner face of the otolith. The scale patch on the cheek is
remarkable for its uniform width with six vertical rows of
scales on the upper part and five on the lower part. The
otolith is characterised by a poor distinction ot ostium
and cauda and the small size of the cauda as found in
A. crassiceps. Geographically, this record is just outside
the distribution range of other species of Alionematichthys.
We have refrained from establishing a new species due
to the lack of a male specimen for investigation of the
pseudoclasper morphology.

Fig. 28. Alionematichthys sp. I, A, lateral view of head, AMS 1. 19108-032, male, 77 mm SL; B, ventral view of head, AMS 1.19108-032,
male, 77 mm SL; C, view of left pseudoclaspers from inside, AMS 1.19108-032, male, 77 mm SL; D. inclined lateral view of male copulatory
organ, AMS 1.19108-032, male, 77 mm SL; E, median view of right otolith, WAM P.30305-036, female, 87 mm SL.

127

W. Schwarzhans and P. R. Moller

Fig. 29. Alionematichthys sp.2, WAM P.251 1 1-047, female, 83 mm SL, A, lateral view of head, B, ventral view of head; C, median view of
right otolith.

Dinematichthys Bleeker, 1855

Type species: Dinematichthys iluoecoeteoides Bleeker,
1855 (type locality Kepulauan Batu, Sumatera Barat
Province, Indonesia). Gender masculine.

Dinematichthys Bleeker, 1855 in part. — Cohen and
Nielsen 1978; Cohen and Hutchins 1982; Machida 1994;
Nielsen et al. 1999.

Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed high on
snout at about 1/2-1/2.5 distance from lip to anterior rim
of eye; head completely and continuously covered with
scales on cheeks, opercle and occiput; cirri on snout; male
copulatory organ with two pairs of small pseudoclaspcrs;
outer pseudoclasper a simple flap, inner pseudoclasper a
flap of about half size of outer, with two or three lobes;
size up to about 110 mm SL; prccaudal vertebrae usually
11 (10-12), total vertebrae 41-45, dorsal fin rays 75-92,
anal fin rays 59-71, V in D 2.1-2.4; upper preopercular pore
absent; sulcus of otolith with ostium at least two times as
long as cauda; maxilla expanded postero-ventrally.

Comparison. Dinematichthys is readily recognised
by its uniform and continuous head squamation including
cheeks, opercle and occiput, usually visible without removal
of mucus. The only other genus sharing this character is
Porocephalichthys gen. nov., which does not overlap in
distribution with Dinematichthys and which differs in
many other characters (see Porocephalichthys gen. nov.).
Even when in doubt, the combination of the absence of the
upper preopercular pore, the small and simple shaped inner
and outer pseudoclaspers, and the otolith with the sulcus
divided into ostium and cauda, usually allow a reliable
identification.

Species. Dinematichthys contains two species, the
widely distributed D. iluocoeteoides known throughout
the Indo-west Pacific and D. trilobatus sp. nov., endemic
to the isolated Christmas and Cocos Islands of the Indian
Ocean.

Remarks. The holotype was originally part of Bleeker’s
collection, who described the first ever dinematichthyine
in 1855, Dinematichthys iluocoeteoides, from a single

128

Dinematichthyine fishes of the Indo-west Pacific, Part IV

specimen from Batu Island (now Kcpulauan Batu, Sumatra
Barat Province), off western Sumatra, Indonesia.

Bleeker’s holotype has been lost (see extensive
discussion in Cohen and Nielsen (1978)). A redefinition of
the genus Dinematichthys and the species D. iluocoeteoides
is therefore necessary.

Cohen and Nielsen (1978) discussed the nature of a
specimen from Bleeker’s original collection (assigned by
Gunther (1862)) kept in London (BMNH 1868.2.28.65), but
concluded that it could not be the original type (though it
was later referred to as the holotype by Eschmeyer (1998)
(also Online Catalog of Fishes, version 19 Sept. 2008,
http://www.calacademy.org/research/ichthyology/catalog/
fishcatmain.asp). It is a male specimen and, although
dried and shrivelled, is here designated as neotype since it
resembles Bleeker’s original description in all important
aspects, and comes from the type locality and Bleeker’s
original collection.

Dinematichthys iluocoeteoides Bleeker, 1855

(Figs 30-32, Tables 1, 13)

Dinematichthys iluocoeteoides Bleeker, 1855: 319;
Gunther 1862; Wounnes and Bayne 1973: 32; Cohen and
Nielsen 1978:58; Dor 1984: 59; Winterbottom etal. 1989:
14; Nielsen et al. 1999: 130.

Dinematichthys indicus Machida, 1994:451; Nielsen et
al. 1999: 130; Schwarzhans and Moller2005: 78.

Dinematichthys randalli Machida, 1994: 456; Nielsen
etal. 1999: 130.

Material examined. (1207 specimens, 11-112 mm
SL). NEOTYPE - BMNH 1862.2.28.65, male, 61 mm
SL, Batu Island, Indonesia, purchased from P. Bleeker.
PARATYPES of D. indicus : ROM 37813-2, 3 males 45,
56 and 92 mm SL, 11 females, 49-77 mm SL, Chagos
Archipelago, Indian Ocean, 7°17’30”N, 7°23’56”E, depth
1-2 m, ROM 58269, 5 males, 48-81 mm SL, 3 females,
36-64 mm SL. 12°23’52”S, 43°30’00”E, Chissioua Dzaha,
Comoros, Indian Ocean.

Additional specimens. AMS I. 17491-006, 1 male, 56
mm SL, 1 female, 59 mm SL, 9°09’S, 159°48’E, Savo
Island, Solomon Islands, 0-10 m; AMS 1. 17492-015, 2
females, 41-48 mm SL, 09°0’S, 160°06’E, Florida Island,
Solomon Islands; AMS I. 20775-092, 1 male, 69 mm SL,
Raine Island, Great Barrier Reef, Queensland, Australia;
AMS I. 22612-010, 1 male 56 mm SL and 2 females, 48
and 55 mm SL, 15°49’S, 145°50’E, Escape Reef, Great
Barrier Reef, Queensland, Australia; AMS I. 33693-008, 1
male, 66 mm SL, 11°42’45”S, 144°04’E, Great Detached
Reef, Queensland, Australia, 6-19 m, AMS I. 33708-062,
1 female, 50 mm SL, 10°59’98”S, 144°01’22”E, Reef
10-418, Queensland, Australia, 2-9 m, AMS I. 37936-070,

1 female, 71 mm SL, 15°00’43”S, 168°03’30”E, Maewo
Island, Vanuatu; BMNH 2000.4.19.370-382, 12 specimens,
57-103 mm SL, Abu Dhabi, Persian Gulf; BPBM 8023, 1
male, 86 mm SL, Marshall Islands; BPBM 8072,2 females,
49-65 mm SL, Palau; BPBM 40933, 2 females, 36 and 59

mm SL, Chuuk, Caroline Islands, 0-4 m; BPBM 10816, 1
female, 59 mm SL, Guadalcanal, Solomon Isis; BPBM
11358, 1 male, 107 mm SL and 1 female, 93 mm SL, Fiji;
BPBM 40935, 1 male, 47 mm SL, Ryukyu Islands, Japan;
BPBM 17704, 1 female, 53 mm SL, Caroline Islands;
BPBM 21795, 1 female, 72 mm SL, Mauritius; BPBM
40937, 1 female, 55 mm SL, Similan Island, Thailand;
BPBM 40939, 2 females, 32 and 47 mm SL, Seychelles;
BPBM 28591, 1 female, 73 mm SL, Philippines; BPBM
29149, 2 females, 55 and 67 mm SL, Marshall Islands;
BPBM 30491, I female, Flores, Indonesia; BPBM 30860,
1 male, 49 mm SL, Persian Gulf; BPBM 30861, 1 female,
54 mm SL, Persian Gulf; BPBM 35199, 1 female, 80 mm
SL, 27° 04’N, 142°12’30”E, Chichi-jima, Ogasawara
Islands, Japan; BPBM 35276,1 male, 74 mm SL, 1 juvenile.

Table 13. Meristic and morphometric characters of Dinematichthys
iluocoeteoides Bleeker, 1855.

Neotype +

Neotype

BMNH

578 non-types

1862.2.28.65

Mean (range)

Standard length in mm
Meristic characters

55.7(11-112)

579

Dorsal fin rays

83.1 (75-92)

137

Caudal fin rays

16.0(15-16)

Anal fin rays

65.3(59-71)

141

Pectoral fin rays

22.7 (21-24)

Prccaudal vertebrae

11.0(10-12)

151

Caudal vertebrae

31.5(30-34)

153

Total vertebrae

42.6(41-45)

153

Rakers on anterior gill arch

15.4(13-19)

Pseudobranchial filaments

D/V

6.0 (5-7)

145

D/A

22.7(19-26)

144

V/A

13.1 (12-14)

145

Morphometric characters in % of SL

Head length

26.2

26.4(25.8-27.1)

Head width

15.2

15.1 (12.2-20.9)

Head height

11.4

16.7(11.4-18.9)

Snout length

4.9

5.9 (4.9-6.5)

Upper jaw length

13.5

13.4(12.7-13.8)

Diameter of pigmented eye

2.8

2.7 (2.2-3.3)

Diameter of pupil

1.2

1.6 (1.2-2.0)

Interorbital width

4.9

6.5 (4.9-7.5)

Posterior maxilla height

6.2

4.6 (4.1-6.2)

Postorbital length

18.8

18.6(17.8-19.2)

Preanal length

45.7

47.2 (44.3-50.6)

Predorsal length

31.6

30.8(29.2-31.8)

Body depth at origin of anal
fin

19.4

19.3(16.3-21.4)

Pectoral fin length

10.5

14.6(10.5-16.2)

Pectoral fin base height

6.1

6.0 (5.2-6.9)

Ventral fin length

17.7

23.5(17.7-25.6)

Base ventral fin - anal fin
origin

27.4

27.1 (24.1-37.0)

129

W. Schwarzhans and P. R. Moller

Fig. 30. Sample sites of O Dinematichthys iluocoeteoides Bleeker, 1855, O* holotype of D. iluocoeteoides , O* holotype of D. indicus,
O* holotype of D. randalli, © D. trilobatus sp. nov. and © Porocephalichthys dasyrhynchus (Cohen and Hutchins, 1982). One symbol may
represent several samples.

Fig. 31. Dinematichthys iluocoeteoides Bleeker, 1855, A, fresh dead, BPBM 35594, female, 80 mm SL; B, neotype, BMNH 1862.2.28.65,
male, 61 mm SL.

21 mm SL, Ogasawara Islands, Japan; BPBM 35594, 1
female, 80 mm SL, Seychelles; BPBM 38149,3 males, 66,
71 and 102 mm SL and 2 females, 56 and 65 mm SL, Tonga;
CAS 14255,6 males, 41-83 mm SL, 6 females 50-102 mm
SL, 7°14’32”N, 144°27’12”E, Ifalik Atoll, Micronesia, 0-4
m; CAS 35340,13 female, 34-72 mm SL, 10 males, 28-60
mm SL, Maldives; CAS 35341,19 males, 32-66 mm SL,
31 females, 29-66 mm SL, 1 juvenile, 23 mm SL,

Seychelles; CAS 227304, 1 male, 70 mm SL, Madang,
Papua New Guinea; CAS 65665, 4 males, 50-75 mm SL,
1 female, 55 mm SL, Madang, Papua New Guinea; CAS
65669, 1 male, 67 mm SL and 2 females, 53 and 63 mm
SL, Madang, Papua New Guinea; CAS 227289. 1 female,
76 mm SL, Madang, Papua New Guinea; CAS 227281, 1
male, 59 mm SLand 1 female, 64 mm SL, Kapingamarangi
Atoll, Micronesia; CAS 227290, 1 female, 46 mm SL, Viti

130

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Fig. 32. Dinematichthys iluocoeteoides Bleekcr, 1855. A, lateral view of head, BPBM 40937. female, 55 nun SL; B. lateral view of head,
USNM 384592, male, 72 mm SL; C, dorsal view of head, USNM 384592, male, 72 mm SL; D, ventral view of head, USNM 384592, male
72 mm SL; E. lateral view of head, USNM 358342. male, 36 mm SL; F, view of left pseudoclaspers from inside, holotype; G, ventral view of
male copulatory organ, BPBM 35276,74 mm SL; H, inclined lateral view of male copulatory organ, BPBM 35276, 74 mm SL; I, view of left
pseudoclaspers from inside, BPBM 35276, 74 mm SL; J, view of left pseudoclaspers from inside, USNM 319902, male, 71 mm SL; K, view
of left pseudoclaspers from inside, SMNS 22544, 52 mm SL; L, median view of right otolith, USNM 199680, male, 78 mm SL; M, ventral
view of right otolith, USNM 199680, male, 78 mm SL; N, median view of right otolith, WAM P.31213-007, female, 52 mm SL.

131

W. Schwarzhans and P. R. Moller

Levu north, Fiji; CAS 222534, 2 females, 58 and 63 mm
SL, Fiji; CAS 222535, 2 females, 24 and 48 mm SL, Fiji;
CAS 222539,4 females, 55-68 mm SL, Fiji; CAS 222541,
female 70 mm SL, Fiji; CAS 227292, 1 female 65 mm SL,
Fiji; CAS 222546, 2 males, 41 and 64 mm SL, 1 female,
43 mm SL, Fiji; CAS 222548, 2 males, 52 and 58 mm SL,
4 females, 41-70 mm SL, 4 juveniles, 15-26 mm SL, Fiji;
CAS 222549, I male, 60 mm SL, 4 females, 43-65 mm
SL, Fiji; CAS 222557, 1 male, 74 mm SL, Fiji; CAS
222560, 1 male, 58 mm SL, 1 female, 55 mm SL, Fiji; CAS
222562,2 males, 47 and 76 mm SL, 1 female with embryos,
50 mm SL, Fiji; CAS 222564,1 male, 49 mm SL, Fiji; CAS
222568, 1 female, 45 mm SL, Fiji; CAS 227306, 1 male,
60 mm SL, Fiji; CAS 227287, 3 males, 38, 45 and 75 mm
SL, 11 female, 40-73 mm SL, 2 juveniles, 19 and 23 mm
SL, Fiji; CAS 222573, 2 females, 58 and 74 mm SL, Fiji;
CAS 222574, 1 female, 38 mm SL, Fiji ; CAS 222575, 1
male, 52 mm SL, 2 females, 41 and 73 mm SL, Fiji; CAS
222577, 3 females, 30-72 mm SL, 2 juveniles, 18 and 22
mm SL, Fiji; CAS 222578, male, 47 mm SL, Fiji; CAS
222579, 1 male, 53 mm SL, Fiji; CAS 222580, 1 female,
50 mm SL, Fiji; CAS 222581, 1 female, 52 mm SL, Fiji;
CAS 222587, 1 male, 78 mm SL, 1 female, 32 mm SL, Fiji;
CAS 222591, female, 86 mm SL, Fiji; CAS 222591, female,
86 mm SL, Fiji; CAS 227308, 2 females, 57 and 59 mm
SL, Fiji; CAS 222593, 2 males, 36-52 mm SL, 3 females,
32-51 mm SL, Fiji; CAS 222597, 8 females, 35-82 mm
SL, Fiji; CAS 222599, 4 males, 48-75 mm SL, 3 females,
65-75 mm SL, Fiji; CAS 222600, 2 males, 78 and 87 mm
SL, 1 female, 78 mm SL, Fiji; CAS 227296, 1 male, 48 mm
SL, Fiji; CAS 222602,4 females, 61-80 mm SL, Fiji; CAS
222603, 1 male, 81 mm SL, 6 females, 65-90 mm SL, Fiji;
CAS 227309, 4 males, 51-83 mm SL, 2 females, 65-73
mm SL, Fiji; CAS 227300, 1 female, 82 mm SL, Fiji; CAS
222609, 2 females, 61 and 80 mm SL, 8 juveniles, 14-33
mm SL, Fiji; CAS 222611, 4 females, 35-72 mm SL, 2
males, 44-86 mm SL, 4 juveniles, 20-25 mm SL, Fiji; CAS
227310, 1 male, 53 mm SL, 1 female, 73 mm SL; CAS
222615, 2 females, 60 and 80 mm SL, Fiji; CAS 222617,
2 females, 49-75 mm SL, 2 males, 44 and 50 mm SL, Fiji;
CAS 222620, 2 females, 39 and 75 mm SL, Fiji; CAS
227302, 2 females, 65 and 75 mm SL, Fiji; CAS 222624,

1 male, 75 mm SL, Fiji; CAS 222627, 1 male, 81 mm SL,
Fiji; CAS 222634, 2 males, 59 and 85 mm SL, 1 juvenile,
19 mm SL; CAS 222639, 1 female, 60 mm SL, 1 male, 58
mm SL, Fiji; KAUM-I. 10660, 1 male, 42 mm SL,
Okinoerabu Island, Kagoshima, Japan; MNHN 1965-0431,

2 females, 54 and 70 mm SL, 23°20’0”S, 43°31 ’0”E,
Madagascar; MNHN 1977-0893, 1 female, 42 mm SL,
29°15’0”N, 34°45’0”E, Gulf of Aqaba, Israel; MNHN
1980-0243, 1 female, 90 mm SL, 22°26’0”S, 166°26’0”E,
New Caledonia; MNHN 1980-0563, I male, 112 mm SL,
17°30’0”S, 167°30’0”E, Pleiades Nord, Loyalty Islands,
15-17 m; EX MNF1N 1980-0851, 1 male, 100 mm SL, 1
female, 66 mm SL, 17°30’0“S, 167°30’0”E, lie Solitaire,
ofTNoumea, New Caledonia; NTM S. 13676-031,2 females.

58 and 70 mm SL, 05°10’S, 145°50’3”E, Tab Island,
Madang, Papua New Guinea, 15-24 m; ROM 37811, 3
males, 50, 50 and 60 mm SL, 05°25’2r’S, 071°46’52”E,
Isle du Coin, Chagos Archipelago; ROM 37812,22 males,
30-81 mm SL, 18 females, 33-68 mm SL, 3 juveniles,
20-24 mm SL, 05°25 W’S, 071 °46’00”E, Isle du Anglaise,
Chagos Archipelago; ROM 42314,1 female, 42 mm SL, 1
male, 48 mm SL, 09°28’00”S, 159°49 ! 00”E, Honiara,
Solomon Islands; ROM 50299, female, 42 mm SL, male
40 mm SL, 09°28’00”S, 159°42’00”E, 10 km W of Honiara,
Solomon Islands; ROM 55144, 63 male, 09°12’16”S,
123°27’15”E, Tonga Point. Philippines; ROM 58267, 6
females, 32-53 mm SL, 5 males, 32-58 mm SL, 2 juveniles,
25 and 25 mm SL, 12°23’52'’S, 43°30’00”E, Point
Chongochahari, Comoros; ROM 68122,4 females, 28-65
mm SL, 3 juveniles, 12-22 mm SL, 12°23’52”S,
43°30'00”E, Chissioua Dzaha, Comoros; SAIAB 35108,
male, 63 mm SL, female, 54 mm SL, 22°00'N, 120°45’E,
Wanlitong, Taiwan; SAIAB 53381, 2 females, 27 and 41
mm SL, 16°18’N. 119°54’E, Luzon, Bolinao, Philippine
Islands; SMNS 13691, 2 females, 61 and 72 mm SL,
27°4I’30”N, 34°08’10”E, Ras Mohammed, Red Sea,
0.5-4.5 m; SMNS 17126,1 female, 28 mm, SL, 19°40’23”S,
63°25’58”E, Rodrigues Island, Mascarcne Islands, 0.3-0.9
m; SMNS 17206,4 females, 40-48 mm SL, 3 males, 39-87
mm SL, 19°40’25”S, 63°25’59’'E, Rodrigues Island,
Mascarene Islands, 0-2 m; SMNS 18674, 2 males, 59-62
mm SL, 1 female, 80 mm SL, 08°22’06”S, 116°04’46”E,
Lombok, Indonesia, 1.5-3.8 m; SMNS 22544, 1 male, 54
mm SL, 29°09'44”S, 34°41’34”E, Gulf of Aqaba, Red
Sea, 0-3 m; TAU 11692, 2 females, 46 and 51 mm SL, 4
males, 42-56 mm SL, Seychelles; TAU 11808, I male, 70
mm SL, I female, 52 mm SL, Seychelles; TAU 11830, 3
specimens, Seychelles; UF 173089, 1 females, 81 mm SL,
1 male, 55 mm SL, Enewetak Atoll, Marshall Islands;
USNM 376199, 1 female, 53 mm SL, Buru Island,
Indonesia; USNM 99066, 1 female, 55 mm SL, Cagayanes,
Philippines; USNM 99223, 1 female, 34 mm SL, Ragay
Gulf, Philippines; USNM 99226, 1 male, 63 mm SL,
Dalaganem, Philippines; USNM 133865, 1 male, 80 mm
SL, Tahiti; USNM 142011, 2 females, 62-68 mm SL,
Rongelap Atoll, Marshall Islands; USNM 142016, 1 male,
82 mm SL, Bikini Atoll, Marshall Islands; USNM 374223,

1 male and 1 female, 46-85 mm SL. Bikini Atoll, Marshall
Islands; USNM 142019, 4 specimens, 89-108 mm SL,
Bikini Atoll, Marshall Islands; USNM 374224, 1 female,
91 mm SL, Bikini Atoll, Marshall Islands; USNM 166807,

2 males and 2 females, 60-82 mm SL, Amo Atoll, Marshall
Islands; USNM 376193,3 females, 52-80 mm SL, Onotoa,
Gilbert Islands, Kiribati; USNM 199680,5 females, 52-65
mm SL, 5 males, 40-83 mm SL, 12°10’S, 44°23’E,
Comoros; USNM 206304, 25 specimens, 34-83mm SL,
Saint Anne Island, Seychelles; USNM 201252, 1 male, 42
mm SL, Ambon, Maluku Islands, Indonesia; USNM
394973,3 females, 31-71 mm SL, Tutuila, Samoa; USNM
223324,2 specimens, 22- 38 mm SL, Pohnpei, Micronesia;

132

Dinematichthyine fishes of the Indo-west Pacific, Part IV

USNM 223412, 6 specimens, 37-62 mm SL, Pohnpei,
Micronesia; USNM 223426, 1 female, 63 mm SL, Pohnpei,
Micronesia; USNM 223508, 14 specimens, 25-81 mm SL,
Pohnpei, Micronesia; USNM 223558,3 specimens, 60-80
mm SL, Pohnpei, Micronesia; USNM 224330,5 specimens,
28-75 mm SL, Pohnpei, Micronesia; USNM 224331, 5
specimens, 23-77 mm SL, Pohnpei, Micronesia; USNM
244012, 1 male, 77 mm SL, 1 female, 32mm SL, 17°45’S,
177°04’W, Malolo Island, Fiji; USNM 244013, 1 male and
4 females, 24-63 mm SL, Malolo Island, Fiji; USNM
263660, 21 specimens, 20-75 mm SL, Lemus, Kavieng,
Papua New Guinea; USNM 263667, 5 specimens, 30-65
mm SL, Red Sea; USNM 263668, 1 female, 50 mm SL,
Marshall Islands; USNM 263673,7 specimens, 40-86 mm
SL, 16°21’S, 43°59’E, Madagascar; USNM 377198, 15
specimens, 45-83 mm SL, 01°33’S, 144°59’E, Papua New
Guinea, 0-15 m; USNM 377209, 1 male and 1 female,
53-64 mm SL, 10°52’N, 120°56’E, Palawan, Philippines,
0-14 m; USNM 377191, 1 female, 58 mm SL, 08°51’N,
123°24’E, Zamboanga, Philippines; USNM 263692, 15
specimens, 51-80 mm SL, 09°10’N, I23°26’E, Negros,
Philippines, 0-3 m; USNM 377200,3 males and 4 females,
38-82 mm SL, Weligama, Sri Lanka; USNM 263708, 1
specimen, 56 mm SL, Vuro Island, Fiji; USNM 263712,
27 specimens, 27-75 mm SL, 10°30’S, 44°21 ’E, Comores;
USNM 263750, 12 specimens, 22-71 mm SL, Kiriwina,
Papua New Guinea; USNM 263756, 18 specimens, 39-83
mm SL, Bougainville, Solomon Islands; USNM 267192,
4 specimens, 17-58 mm SL, Seychelles; USNM 394974,
4 males, 27-36 mm SL, 2 females, 42 and 43 mm SL, Mahe,
Seychelles; USNM 300092, 2 males and 1 female, 44-69
mm SL, 20°24’N, 121°55’E, Batanes, Philippines; USNM
300094,1 female, 21 °07’N, 121 °56’E, Batanes, Philippines;
USNM 300100, 1 female, 20°17’N, 121°50’E, Batanes,
Philippines; USNM 300103, I male, 50 mm SL, 22°54’N,
121°54’E, Batanes, Philippines; USNM 374228, 1 male,
57 mm SL,04°52’N, 119°26’E, Sibutu, Philippines; USNM
319898, 1 female, 78 mm SL, 20°34’S 166°14’E, Loyalty
Islands, New Caledonia, 0-5 m; USNM 319902, 2 males
50 and 71 mm SL, 5 females, 20°34’S, I66°14'E, Loyalty
Islands, New Caledonia; USNM 334123, 20 specimens,
26-89 mm SL, Tongatapu, Tonga; USNM 334125, 9
specimens, 28-75 mm SL, Tongatapu, Tonga; USNM
334126, 2 specimens, 23-93, mm SL, Tongatapu, Tonga;
USNM 334127, 13 specimens, Tongatapu, Tonga; USNM
374209,11 specimens, 35-79 mm SL, 21 °20’S, 174°58’W,
Tonga; USNM 336510, 9 specimens, 19°16’S, 174°22’W,
Tonga; USNM 338465, 4 females, 31-98 mm SL, Vava’u
Group, Tonga; USNM 394977, 6 specimens, 54-67 mm
SL, 18°44'31”S, 174°06’36”W, Vava’u Group, Tonga;
USNM 377199, 5 specimens, 42-75 mm SL, 10°35’N,
122°08’E, Panay, Philippines; USNM 376156, 1 female,
47 mm SL, 10°28’N, 122°28’E, Guimaraes, Philippines;
USNM 394983, 5 females, 43-72 mm SL, 1 male, 58 mm
SL, 16°47’13”S, 168°2r36”E, Epi, Vanuatu, 1-10 m;
USNM 352643, 6 specimens, 52-55 mm SL, Ryukyu

Islands; USNM 355820, 1 male and I female, 40-66 mm
SL, Papua New Guinea; USNM 357210,1 juvenile, 23 mm
SL, 10°40’S 165°47’E, Solomon Islands; USNM 358342,
1 male, 36 mm SL, 10°16’S, 166°18’E, Solomon Islands;
USNM 384592, 1 female, 63 mm SL, 1 male, 72 mm SL,
Vanuatu, Efate; USNM 363675, 1 female, 79 mm SL,
Vanuatu; USNM 363908, 2 females, 60-88 mm SL,
15°00’S, 168°03’E, Maewo, Vanuatu; USNM 365322, 1
male and I female, 80-102 mm SL, 08°23’S, 162°5UE,
Stewart Island, Solomon Islands; USNM 365613, 4
specimens, 42-57 mm SL, 13 o 21’30”S, 176°10’10”W,
Wallis Island, Wallis and Futuna Islands, 1-13 m; USNM
377236, 2 males, 41-80 mm SL, Bugabag, Papua New
Guinea; USNM 365823, 4 specimens, 42-75 mm SL,
Hermit Island, Papua New Guinea; USNM 365826, 7
specimens, 58-66 mm SL, Hermit Island, Papua New
Guinea; USNM 365829, 1 female, 37 mm SL, Massas,
Papua New Guinea; USNM 366216, 6 specimens, 51-83
mm SL, Chagos Archipelago; USNM 366227, 3 females,
25-50 mm SL, 27°16’N, 33°47’E, Egypt; USNM 366228,
1 male, 45 mm SL, 26°08’N, 34°16’E, Egypt; USNM
366229, 1 female, 68 mm SL, 27°18’N, 33°47'E, Egypt;
USNM 366230, 1 male and 1 female, 40-60 mm SL,
Mauritius; USNM 366231, 2 juveniles, 17-26 mm SL,
16°25’S, 59°36’E, Cargados Carajos, Mascarene Islands;
USNM 366232, 3 females, 38-68 mm SL, 16°43’S,
59°35’E, Cargados Carajos, Mascarene Islands; USNM
366233, 7 specimens, 35-65 mm SL, 16°27’S, 59°36’E,
Cargados Carajos, Mascarene Islands; USNM 366234, 9
males and 24 females, 31-70 mm SL, 10°I9’S, 56°35’E,
Agalega, Mauritius; USNM 366457,2 specimens, Agalega,
Mauritius; USNM 366459, 8 males and 4 females, 32-86
mm SL, 07°15’S, 72°22’E, Chagos Archipelago; USNM
366460, I male, 54 mm SL, 07°15’S, 72°22’E, Chagos
Archipelago; USNM 366461,5 males and 8 females, 42-84
mm SL, 07°15’S, 72°22’E, Chagos Archipelago; USNM
366462, 1 female, 45 mm SL, 13°49’S, 72°24’E, Chagos
Archipelago; USNM 366463, 2 females, 57-60 mm SL,
07°14’S,72°23’E, Chagos Archipelago; USNM 366464,3
males and 2 females, 39-69 mm SL, 07°20’S, 72°27’E,
Chagos Archipelago; USNM 366465,5 males and 1 female,
40-59 mm SL, Marsa Mokrakh, Egypt; USNM 366466,4
females, 27—41 mm SL, Egypt; USNM 366467, 1 female,
68 mm SL, Hikkaduwa, Sri Lanka; USNM 366468, 6
specimens, 40-82 mm SL, Hikkaduwa, Sri Lanka; USNM
366469,1 male, 72 mm SL, Hikkaduwa, Sri Lanka; USNM
366472, 33 specimens, 22-92 mm SL, Egypt; USNM
376219, 6 males and 1 female, 21-65 mm SL, 05°52’S,
112°37’E, Java, Bawaean Island; USNM 366476, 47
specimens, 16°28’S, 59°40’E, Cargados Carajos, Mascarene
Islands; USNM 366477, I male, 71 mm SL, Hikkaduwa,
Sri Lanka; USNM 366478, 1 male and 1 female, 71-84
mm SL, Galle, Sri Lanka; USNM 366479, 1 female, 60
mm SL, Hikkaduwa, Sri Lanka; USNM 366483, 1 female,
58 mm SL. 07°14’S, 72°23’E, Chagos Archipelago; USNM
366484, 2 females, 42-57 mm SL, 07°20’S, 72°27’E,

133

W. Schwarzhans and P. R. Muller

Chagos Archipelago; USNM 366485, 2 males and 6
females, 44-67 mm SL, 07°13’S, 72°25’E, Chagos
Archipelago; USNM 366486, 1 female, 84 mm SL,
07°14’S, 72°23’E, Chagos Archipelago; USNM 366487, 1
male and 4 females, 44-69 mm SL, 07°16’S, 72°28’E,
Chagos Archipelago; USNM 366491, 1 female, 41 mm SL,
02°50’N, 95°56’E, Sumatra; USNM 366492, 3 males,
35-67 mm SL, 04°14’S, 152°10’E, Bismarck Archipelago,
Papua New Guinea; USNM 376220, 1 male, 65 mm SL,
03°47’S, 128°06’E, Maluku, Indonesia; USNM 376200, 1
female, 39 mm SL, 05°52’S, 110°25’E, Karimundjawa,
Java, Indonesia; USNM 366500, 1 male and 7 females,
39-75 mm SL, 05°51 ’S, 106°34’E, Pulau Seribu, Indonesia;
USNM 366501, 19 specimens, 16-48 mm SL, 01°33’S,
144°59’E, Papua New Guinea; USNM 376205,5 specimens,
48-74 mm SL, Madang, Papua New Guinea; USNM
366504,3 specimens, Madang, Papua New Guinea; USNM
366506, 1 male and 3 females, 68-87 mm SL, Briwadi
Island, Kiriwina Islands (Trobriand), Papua New Guinea;
USNM 376162, 31 specimens, 07°15’S, 72°22’E, Chagos
Archipelago; USNM 366513,1 juvenile, 29 mm SL, Buroa,
Egypt; USNM 366514, 1 female, 61 mm SL, 15°32’N,
40°00’E, Ethiopia; USNM 366515, 14 specimens, 15-67
mm SL, Gulf of Aqaba, Egypt; USNM 366516, 1 female,
74 mm SL, Buroa, Egypt; USNM 366517, 2 males and 1
female, 43-71 mm SL, Sinai, Egypt; USNM 366519, 4
females, 41-58 mm SL, 16°28'S, 59°37’E, Cargados
Carajos, Mascarcne Islands; USNM 366520,15 specimens,
34-67 mm SL, North Island, Agalega, Mauritius; USNM
376201, lOspecimens, 34-69 mm SL,05°10’S, 145°5UE,
Papua New Guinea; USNM 366523, 1 female, 61 mm SL,
Madang, Papua New Guinea; USNM 376191,1 female, 45
mm SL, Madang, Papua New Guinea; USNM 366526, 13
specimens, 39-61 mm SL, Kiriwina Islands (Trobriand),
Papua New Guinea; USNM 366527, 11 specimens, 59-91
mm SL, 12°46’N, 44°59’E, Aden; USNM 366529, 3
specimens, 12°53’S, 45°16’E, Comores; USNM 366534,
4 specimens, 07°56’N, 81 °34’E, Sri Lanka; USNM 366535,
21 specimens, 26-75 mm SL, 05°24’S, 53° 13’E, Comores;
USNM 366551,5 specimens, 21-41 mm SL, Israel; USNM
366552, 22 specimens, 11-98 mm SL, Gulf of Aqaba,
Egypt; USNM 366553, 13 specimens, 51-72 mm SL,
15°30’N, 39°54’E, Ethiopia; USNM 366571, 1 male, 54
mm SL, Guadalcanal, Solomon Islands; USNM 366573, 1
male, 73 mm SL, New Georgia, Solomon Islands, USNM
366590, 20 specimens, 30-80 mm SL, 16°36’S, 59°31’E,
Cargados Carajos, Mascarene Islands; USNM 366591, 31
specimesn, 28-75 mm SL, 16°26’S, 59°36’E, Cargados
Carajos, Mascarene Islands; USNM 366593,30 specimens,
28-74 mm SL, 16°25’S, 59°36’E, Cargados Carajos,
Mascarene Islands; USNM 366594, 5 specimens, 35.81
mm SL, 16°15’S, 59°33’E, Cargados Carajos, Mascarene
Islands; USNM 374229, 1 female, 51 mm SL, 09°13’N,
123°28’E, Negros, Philippines; USNM 374225, 1 male, 66
mm SL, 09°31’N, 123°40’E, Philippines; USNM 366679,
1 female, 86 mm SL, Kwajalein Atoll, Marshall Islands;

USNM 366692, 2 females, 68-72 mm SL, 21°55’N,
120°50’E, Taiwan; USNM 366693, 2 specimens, 48-64
mm SL, 21°55’N, 120°48’E, Taiwan; USNM 366697, 20
specimens, 37-74 mm SL, Red Sea; USNM 366701, 1
specimen. 47 mm SL, 09°03’N, 123°07’E, Philippines;
USNM 366702, 4 specimens, 27-43 mm SL, Bararin,
Philippines; USNM 366704, 1 male, 49 mm SL, Tagauyan,
Philippines; USNM 366720, 1 male, 55 mm SL, Balabac,
Philippines; USNM 366721. I female, 44 mm SL, 11°42’S,
167°51 ’E, Vanikolo Islands, Santa Cruz Islands, Solomon
Islands,; USNM 366839, 7 specimens, 59-80 mm SL,
18°08’S, 178°24’E, Fiji; USNM 366840, 1 male, 78 mm
SL, 19°09’S, 179°45’E, Fiji; USNM 366842,6 specimens,
45-77 mm SL, 18°58’S, 179°52’W, Fiji; USNM 366844,
4 specimens, 55-76 mm SL, 21°38’S, 178°45'W, Fiji;
USNM 366845, 6 males and 1 female, 44-81 mm SL,
18°55’S, 178°33’W, Fiji; USNM 366846, 6 specimens,
45-75 mm SL, 18°57'S, 178° 17’E, Fiji; USNM 376206, 6
specimens, 40-111 mm SL, 17°06’S, 177° 13’E, Fiji;
USNM 374208, 3 females, 32-60 mm SL, 13°23’S,
176° 11 ’ W, Wallis Island, Wallis and Futuna Islands; USNM
263698, 1 female, 57 mm SL, 01°33’S, 144°59’E, Papua
New Guinea; WAM P.27469-005, 5 males, 65-70 mm SL,
15°50’S, 145°50’E, Escape Reef, Queensland, Australia;
WAM P.27663-003, 2 males 65 and 65 mm SL, 1 female,
35 mm SL, NW Australia; WAM P.27825-031,2 males, 35
and 42 mm SL, 2 females, 39 and 44 mm SL, Bismarck
Sea, Papua New Guinea; WAM P.28191-005, 1 male, 52
mm SL, Coral Sea; WAM P.29627-048, 2 females, Coral
Sea; WAM P.29642-010, 2 females, 42-53 mm SL, Coral
Sea; WAM P.30618-002, 1 male and 3 females, 67-79 mm
SL, Papua New Guinea; WAM P.30633-023, 1 male, 58
mm SL, Papua New Guinea; WAM P.30719-036,1 female,
42 mm SL, Flores Sea, Indonesia; WAM P.30791-002, 1
female, 50 mm SL, New Caledonia; WAM P.30844-050, 3
specimens, Timor Sea; WAM P.31213-058. 1 female, 52
mm SL, Papua New Guinea; WAM P.31437-043, 1 male,
60 mm SL, Timor Sea; YCM-P 34028, 1 male, 54 mm SL,
Kakeroma Island, northern Ryukyu Islands, Japan; ZMUC
P 771654-56, 2 males, 66 and 85 mm SL and 1 female, 77
mm SL, Seychelles; ZMUC P 771657-58, 1 male, 61 mm
SL, and 1 female, 56 mm SL, Mauritius.

Diagnosis. Vertebrae 10-12+30-34=41-45, dorsal
fin rays 75-92, anal fin rays 59-71; eyes large (2.2-3.3%
SL); body moderately slender, snout with many cirri; head
continuously covered with scales, ventrally terminating
behind maxilla; upper preopercular pore absent; outer
pseudoclasper broad-based, small, not extending beyond
hood in resting position; inner pseudoclasper small, with
two lobes; otolith length to height 2.1-2.3 with regularly
curved dorsal rim, otolith length to sulcus length 1.6—1.7,
ostium length to cauda length 3.8^4.8.

Description (Figs 31, 32).The principal meristic and
morphometric characters are shown in Table 13. Mature at
about 45 mm SL. Body slender, with moderately pointed
head profile. Many cirri on snout. Eye size 2.2-3.3 (2.8)%

134

Dinematichthyine fishes of the Indo-west Pacific, Part IV

SL. Head continuously covered with scales on cheeks,
opercle and occiput, ventrally terminating behind maxilla.
Horizontal diameter of scales on body of 85 mm SL male
(ZMUC P771656) about 2.0% SL, in about 28 horizontal
rows. Maxillary ending far behind eye, dorsal margin of
maxillary covered by upper lip dermal lobe, posterior
end expanded with little knob. Anterior nostril positioned
high, 1/2—1/2.5 distance from upper lip to anterior margin
of eye. Posterior nostril small, about one-fifth to one-sixth
the size of eye. Opercular spine with free tip, pointed.
Anterior gill arch with 13-19 (15) rakers, 3 elongate in a
row. Pseudobranchial filaments 2.

Head sensory pores (Fig. 32A-E). Supraorbital pores
2-3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third the size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior):
first anterior pore large, tubular, with cirrus anteriorly.
Preopercular pores: 3 lower, first and second with joined
opening, covered by dermal flap in lateral view, third
tubular; no upper preopercular pore. [See description of
Alionematichthys ceylonensis for position of pores.]

Dentition (of holotype). Premaxilla with 7 outer rows of
granular teeth and 2 inner rows of larger teeth. Anteriormost
teeth in inner row up to 3/4 diameter of pupil. Vomer
horseshoe-shaped, with 3 outer rows of small teeth and 1
inner row of larger teeth up to 2/3 diameter of pupil. Palatine
with 2 outer rows of small teeth and 1 inner row of larger
teeth up to 1 /2 diameter of pupil. Dentary with 5 outer rows
of granular teeth and 1 inner row of larger teeth, up to about
size of diameter of pupil.

Otolith (Fig. 32L-N). Elongate in shape, length to
height 2.1-2.3 and moderately thin (otolith height to otolith
thickness about 2.5). Anterior tip rounded, posterior tip
slightly pointed, expanded. Dorsal rim shallow, gently
curved. Inner face convex, outer face slightly concave to
flat, both smooth. Otolith length to sulcus length 1.6-1.7.
Sulcus medianly positioned, slightly inclined, with
separated colliculi and small notch at ventral rim of sulcus
at joint of ostium with cauda. Ostium length to cauda length
3.8^1.8. Ventral furrow weak, moderately close to ventral
rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined and
5-8 depressed. Parapophyses present from vertebrae 6
to 11 (10-12). Pleural ribs on vertebrae 2 to 11 (10-12).
First anal fin pterygiophore not longer than subsequent
pterygiophore.

Male copulatory organ (Fig. 32G-K). Two pairs of
small pseudoclaspcrs, outer not extending beyond hood in
resting position. Outer pseudoclasper broad based, simple
flap-shaped; inner pseudoclasper about half length of outer
pseudoclasper, with two lobes. Isthmus moderately narrow;
penis curved, slightly longer than outer pseudoclaspers.

Colour. Live colour reported as to yellow or bright to
red (Fig. 31 A). Preserved colour mostly dark brown.

Variability and ontogeny. Despite the wide distribution
of D. iluocoeteoides there are few, if any, regional

variations to be observed. For instance, in life, colouration
of specimens from the Western Indian Ocean is usually
described as bright red while a record from the most north¬
eastern location at the Japanese Bonin Islands was reported
as yellow (personal communication by J. E. Randall).
Another character showing some variation is the amount
of cirri on the snout, which appears to be denser in western
Indian Ocean specimens than in western Pacific specimens.
Specimens from furthest to the southwest (Vanuatu, Fiji,
Tonga) are special for their generally somewhat higher
meristic counts, both in vertebrae (43-45) as well as
dorsal fin rays (84-92). Ontogenetic changes are most
strongly expressed in the squamation of the head, which
is continuous on cheeks, opercle and occiput in adults and
subadults from about 40 mm SL. Smallerjuvenile and larval
specimens show some interruption of the scale coverage on
the head, particularly between cheeks and opercle, while
already complete between cheeks and occiput (Fig. 32E).
The density of cirri on the snout also increases with growth,
i.e. is absent in small specimens below 40 mm SL.

Remarks. Machida (1994) described D. indicus from
the Chagos Archipelago and Comores Islands, i.e. from
the central and western Indian Ocean and D. randalli from
Kosrae, Micronesia. He distinguished D. indicus from
D. randalli by the presence of cirri on the top and sides of
the head behind the eye (vs no cirri on head) and by lower
scale row counts (85-100 vs 107-113). In our review of
more than 1000 specimens from an almost continuous
geographical range, the main diagnostic character, i.e.
presence or absence of cirri on the head could not be
verified. It is true that western Pacific specimens appear to
have generally fewer cirri, but we regard this character as
too weak and too irregularly distributed to be of diagnostic
value. Similarly, the scale row count is considered by us as
a not sufficiently reliable character.

Distinction of these two species from D. iluocoeteoides
was based on comparative data from Bleekcr (1855) and
Sedor and Cohen (1987), with D. iluocoeteoides and was
described as having larger eyes (a little over 5 in head
length / eye diameter vs 7.8-9.4) and having a sheathed
maxillary (vs unsheathed). Again, in our evaluation the eye
size differences are well within the variations observed,
while we failed to recognise a difference of sheathed versus
unsheathed maxillary in any of the specimens other than
possibly caused by preservation. In consequence, all three
nominal species arc here regarded as representing a single
valid species for which D. iluocoeteoides is the senior
synonym.

Comparison. Dinematichthys iluocoeteoides is
closely related to D. trilohatus sp. nov. (see below for
differentiation of the two species). The head squamation
continuing on the cheeks, opercle and occiput, which
usually is easily visible without the removal of mucus,
distinguishes D. iluocoeteoides readily from any other co¬
occurring dinematichthyine species. Confusion may only
be possible in the case of juveniles, which show some gaps

135

W. Schwarzhans and P. R. M 0 ller

in head squamation between the cheeks and the opercle
(Fig. 32E). Also, the absence of the upper preopercular
pore and connection of the cheek squamation with the
occiput squamation still allow a secure identification in
most instances.

Distribution. Dinematichthys iluocoeteoides is the
most widely distributed species of all Dinematichthyini.
ranging from the western Indian Ocean (Red Sea, Arabian
Gulf, East Africa) to the Ryukyu and Ogasawara Islands
in the north-western Pacific and Marshall and Kiribati
Islands, Fiji, Tonga and Samoa in the central and south¬
western Pacific (Fig. 30). The reason for its unusually
wide distribution is unknown, but due to the low degree of
regional variation (except for the sympatric D. trilobatus
sp. nov. of the Christmas and Cocos Islands; see below) is
likely to be of recent origin. Despite this wide distribution,
D. iluocoeteoides usually does not occur in large numbers
at most sampled locations, except for some areas in the
western and central Indian Ocean, e.g. the Comores or the
Chagos Archipelago. Dinematichthys iluocoeteoides is also
notable for its strict association with true reef environments
and particularly so with outer reefs and reef cores.

Dinematichthys trilobatus sp. nov.

(Figs 30, 33, 34, Tables 1, 14)

Material examined. (16 specimens, 37-58 mm SL).
HOLOTYPE - WAM P.29927-005, male, 46 mm SL,
12°05’S, 096°53’E, Direction Island, Cocos-Keeling
Islands, 0.1-2.0 m, G. R. Allen, 24 Feb. 1989. PARATYPES
- ANSP 162877, 49 mm SL, I2°06’30”S, 096°49’35”E;
Turk Reef, Cocos-Keeling Islands, 45-49 m, W.F. Smith-
Vaniz and P. L. Colin, 8 March 1974; WAM P.26106-012,
male, 40 mm SL, 7 females, 45-57 mm SL, 10°29’S,
105°40'E, Rhonda Beach, Christmas Island, 10-12 m, G.
R. Allen and R. Steene, 2 June 1978; WAM P.29002-021,
males, 42, and 50 mm SL, female 58 mm SL, 10°25’S,
105°40’E, Kiritimati, 13-26 m, G. R. Allen and R. Steene,
28 June 1986; WAM P.29927-013), 3 females, 37, 37 and
38 mm SL, same data as the holotype.

Diagnosis. Vertebrae 11+31-32=42-43, dorsal fin
rays 80-87, anal fin rays 60-67; eyes large (2.4-3.0%
SL); body slender, snout with few cirri; head continuously
covered with scales, ventrally extending beyond tip of
maxilla forward of third posterior mandibular pore; upper
preopercular pore absent; outer pseudoclasper broad-based,
small, not extending beyond hood in resting position; inner
pseudoclasper small, with three lobes; otolith length to
height 2.1-2.2, with regularly curved dorsal rim, otolith
length to sulcus length 1.7-1.8, ostium length to cauda
length 3.7-4.2.

Description (Figs 33, 34).The principal meristic and
morphometric characters are shown in Table 14. Mature at
about 40 mm SL. Body slender, with pointed head profile.
Few cirri on snout. Eye size 2.4-3.0 (2.8)% SL. Head
continuously covered with scales on cheeks, opercle and
occiput, ventrally extending beyond tip of maxilla forward

Table 14. Meristic and morphometric characters of Dinematichthys
trilobatus sp. nov.

Holotype

WAM

29927-

005

Holotype +

15 paratypes

Mean (range)

Standard length in mm

47.4 (37-58)

Meristic characters

Dorsal fin rays

82.6 (80-87)

Caudal fin rays

15.9(15-16)

Anal fin rays

64.1 (60-67)

Pectoral fin rays

22.4 (21-24)

Precaudal vertebrae

Caudal vertebrae

31.7(31-32)

Total vertebrae

42.7 (42-43)

Rakers on anterior gill arch

15.4(13-17)

Pseudobranchial filaments

D/V

D/A

22.0(19-24)

V/A

Morphometric characters in % of SL

Head length 26.5

27.1 (25.8-28.0)

Head width

15.0

13.7(12.7-16.6)

Head height

17.0

16.5(15.6-17.5)

Snout length

5.3

5.9 (4.9-6.9)

Upper jaw length

13.4

14.0(12.9-15.0)

Diameter of pigmented eye

2.8

2.7 (2.4-3.0)

Diameter of pupil

1.7

1.8(1.5-2.6)

Interorbital width

6.2

6.9 (6.2-7.9)

Posterior maxilla height

4.5

4.5 (4.1-4.8)

Postorbital length

18.8

19.0(17.8-19.8)

Preanal length

49.1

45.8 (42.6-49.1)

Predorsal length

30.1

31.1 (30.1-32.2)

Body depth at origin of anal fin

16.9

17.6(15.7-20.0)

Pectoral fin length

14.6

14.8(12.5-16.4)

Pectoral fin base height

5.9

6.3 (5.8-7.2)

Ventral fin length

23.5

23.6(20.9-26.1)

Base ventral fin - anal fin origin

27.2

25.7 (23.7-27.5)

of third posterior mandibular pore. Horizontal diameter
of scales on body about 1.3% SL, in 27 horizontal rows.
Maxillary' ending far behind eye, dorsal margin of maxillary
covered by upper lip dermal lobe, posterior end expanded
with little knob. Anterior nostril positioned high, 1/2.5
distance from upper lip to anterior margin of eye. Posterior
nostril small, about 1 /6 the size of eye. Opercular spine with
free tip, pointed. Anterior gill arch with 13-17(15) rakers,
3 elongate in a row. Pseudobranchial filaments 2.

Head sensory pores (Fig. 34A-C). Supraorbital pores
3. Infraorbital pores 6 (3 anterior and 3 posterior), three
posterior pores about one-third size of three anterior
pores. Mandibular pores 6 (3 anterior and 3 posterior): first
anterior pore large, tubular. Preopercular pores: 3 lower,
first and second with joined opening, covered by dermal
ilap in lateral view; third tubular; no upper preopercular

136

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Fig. 33. Dinematichthys trilobatus sp. nov., WAM P.29927-005, holotype, male, 46, mm SL.

Fig. 34. Dinematichthys trilobatus sp. nov., A, lateral view of head, WAM P.29002-021, female, 58 mm SL; B, dorsal view of head, WAM
P.26106-012, female, 57 mm SL; C, ventral view of head, WAM P.26106-012, female, 57 mm SL; D, ventral view of male copulatory organ,
WAM P.29002-021, male, 42 mm SL; E, view ofleft pseudoclaspcrs from inside, holotype; F, inclined lateral view of male copulatory organ,
WAM P.29002-021,42 mm SL; G, median view of right otolith, WAM P.29002-021, male, 42 mm SL; H, ventral view ol right otolith, WAM
P.29002-021, male, 42 mm SL.

pore. [See description of Alionematichthys ceylonensis for
position of pores.]

Dentition (of holotype). Premaxilla with 6 outer rows of
granular teeth and 1 inner row of larger teeth. Anterionnost
teeth in inner row up to 1/3 diameter of pupil. Vomer
horseshoe-shaped, with 2 outer rows of small teeth and 1
inner row of larger teeth up to 1/4 diameter of pupil. Palatine

with 1 outer row of small teeth and 1 inner row of larger
teeth up to 1/4 diameter of pupil. Dentary with 3 outer rows
of granular teeth and I inner row of larger teeth, up to about
1/2 diameter of pupil.

Otolith (Fig. 34G, FI). Elongate in shape, length to
height 2.1-2.2 and moderately thin (otolith height to otolith
thickness about 2.3). Anterior tip slightly pointed, posterior

137

W. Schwarzhans and P. R. Muller

tip pointed, expanded. Dorsal rim shallow, gently curved.
Inner face convex, outer face slightly concave to flat, both
smooth. Otolith length to sulcus length 1.7-1.8. Sulcus
medianly positioned, slightly inclined, with separated
colliculi and marked notch at ventral rim of sulcus at
joint of ostium with cauda. Ostium length to cauda length
3.7-4.2. Ventral furrow weak, moderately close to ventral
rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined and
5-8 depressed. Parapophyses present from vertebrae 6 to 11.
Pleural ribs on vertebrae 2 to 11. First anal fin pterygiophore
not longer than subsequent pterygiophore.

Male copulatory organ (Fig. 34D-F). Two pairs of
small pseudoclaspers, the outer not extending beyond
hood in resting position. Outer pseudoclasper broad based,
simple flap-shaped; inner pseudoclasper about half the
length of outer pseudoclasper, with three lobes. Isthmus
moderately wide; penis curved, about as long as outer
pseudoclaspers.

Colour. Live colour unknown. Preserved colour light
brown.

Comparison. Dinematichthys trilobatus obviously is
closely related to D. iluocoeteoides, from which it differs
mainly in the shape of the inner pseudoclaper with three
distinct lobes (vs two), the more expanded head squamation
ventrally extending beyond tip of maxilla forward of third
posterior mandibular pore, and and only few cirri on the
snout (vs many cirri).

Distribution. Dinematichthys trilobatus is endemic to
the Christmas and Cocos-Keeling Islands, where it replaces
the widespread D. iluocoeteoides (Fig. 30). Dinematichthys
trilobatus represents the only dinematichthyine known from
the Christmas and Cocos Islands and its specific separation
from the widespread D. iluocoeteoides indicates a prolonged
separation history for the fauna of these islands, or a perhaps
a colonisation event with subsequent speciation.

Etymology. Named for the inner pseudoclasper with
three lobes: trilobatus (Latin = three lobed). It is an
adjective.

Porocephalichthys gen. nov.

Type species: Dinematichthys dasyrhynchus Cohen
and Hutchins, 1982 (type locality Rottnest Island, Western
Australia). Gender masculine.

Dinematichthys (non Bleeker, 1855) in part. — Cohen
and Nielsen 1978; Cohen and Hutchins 1982; Machida
1994. Nielsen et al. 1999.

Diagnosis. Genus of Dinematichthyini with following
combination of characters: anterior nostril placed high on
snout at about 1/2.5 the distance from lip to anterior rim
of eye; head completely and continuously covered with
scales on cheeks, opercle and occiput; abundant cirri on
snout; male copulatory organ with two pairs of equally
long pseudoclaspers; outer pseudoclasper a simple flap,
inner pseudoclasper inwardly curved stick-like; size up
to 100 mm SL; prccaudal vertebrae 14, total vertebrae

47-49, dorsal fin rays 96-104, caudal fin rays 17-18,
anal fin rays 62-69, V in D 2.3; upper prcopercular pore
present: four additional pairs of supraorbital pores, one
pair above and in front of eyes, a second above and behind
eyes, two more pairs of pores on occiput in prolongation
of upper preopercular pore; one or two additional posterior
mandibular pores; jaws without canine teeth; no elongate
gill rakers; sulcus of otolith with cauda nearly of the length
of ostium.

Comparison. When originally described, the species
was tentatively placed in the genus Dinematichthys. Cohen
and Hutchins (1982, p. 342) wrote: “Differences between
the two are numerous and when more species are described
and specimens of the true D. iluocoeteoides are re-collected
and become available for study, it may be necessary to
establish the Rottnest Island species in a separate genus”.
After having reviewed all Dinematicthyine genera, we are
confident that D. dasyrhynchus should be placed in a separate
genus. Porocephalichthys resembles Dinematichthys in the
uniform and continuous head squamation including cheeks,
opercle and occiput and in having the anterior nostril placed
high on snout. The other characters of the diagnosis, such as
the many additional pores, total lack of elongate gill-rakers,
number of vertebrae, dorsal, caudal and pectoral fin rays
(Table 1), and the proportions of the sulcus of the otolith
and the pseudoclasper pattern, place Porocephalichthys
well apart from Dinematichthys and any of the other
dinematichthyine genera. The correct phylogenetic position
of the genus needs further investigation.

Species. Porocephalichthys is a monospecific genus
with P. dasyrhynchus endemic to SW Australia.

Etymology. Combined from porocephalus (Latin =
head with pores) and ichthys (Greek = fish), referring to the
many pores on the head. The gender is masculine.

Porocephalichthys dasyrhynchus (Cohen and
Hutchins, 1982)

(Figs 30, 35, 36, Tables 1, 15)

Dinematichthys iluocoeteoides. — Mees I960: 18.

Dinematichthys sp. — Hutchins 1979: 93.

Dinematichthys dasyrhynchus Cohen and Hutchins,
1982: 342; Allen 1985: 2268, fig. 49; Paxton et al. 1989:
316; Nielsen et al. 1999: 130.

Material examined (9 specimens, 43-95 mm SL) (all
from Western Australia). Paratypes: AMS 1. 20245-016,

1 male, 64 mm SL, 2 females, 77-94 mm SL, Horseshoe
Reef, Rottnest Island, 12-15 m, B. C. Russell and J. B.
Hutchins; Non-types: WAM P.27950-011, 2 males, 61
and 64 mm SL, 30°18’S, 115°00’E, Jurien Bay, 4-6 m, N.
Sinclair et al.. 9 April 1983; WAM P.27951-007, 1 female,
95 mm SL, 30°18’S, 115°00’E, Jurien Bay, Osprey Inlet,
Western Australia, 2-5 m, J. B. Hutchins et al., 10 April
1983; ZMUC P77716-18,3 females, 43,46 and 89 mm SL,
32°0LS, 115°29’E, Green Island, Rottnest Island, Western
Australia, 10 April 1978.

Diagnosis. See generic diagnosis.

138

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Fig. 35. Porocephalichthys dasyrhynchus (Cohen and Hutchins, 1982), ZMUC P77726, female, 89 mm SL.

Fig. 36. Porocephalichthys dasyrhynchus (Cohen and Hutchins, 1982), A, lateral view of head, WAM P.27950-011, male, 64 mm SL; B, ventral
view of head, WAM P.27950-011, male, 64 mm SL; C, dorsal view of head, WAM P.27950-011, male, 64 mm SL; D, inclined lateral view
of male copulatory organ, WAM P.27950-011, male, 61 mm SL; E, view of left pscudoclaspers from inside, WAM P.27950-011, male, 64
mm SL; F, view of left pseudoelaspers from inside, WAM P.27950-011, male, 61 mm SL; G, median view of right otolith, ZMUC P77726,
female, 89 mm SL; II, anterior view of right otolith, ZMUC P77726, female, 89 mm SL; I, dorsal view of right otolith, ZMUC P77726,
female, 89 mm SL.

139

W. Schwarzhans and P. R. Moller

Table 15. Meristic and morphometric characters of Porocephalichthys
dasyrhynchus (Cohen and Hutchins, 1982).

Holotype

WAM

P.26614-

010*

Holotype +

6 paratypes and

6 non-types

Mean (range)

Standard length in mm

Meristic characters

74.8 (43-102)

Dorsal fin rays

99.4(96-104)

Caudal fin rays

17.3(17-18)

Anal fin rays

66.4 (62-69)

Pectoral fin rays

26.5 (25-28)

Precaudal vertebrae

Caudal vertebrae

34.1 (33-35)

Total vertebrae

48.1 (47-49)

Rakers on anterior gill arch

21.3(19-26)

Pseudobranchial filaments

D/V

D/A

39.2 (37-43)

V/A

Morphometric characters in % of SL

16.8(16-17)

Head length

26.1

26.4(24.6-28.1)

Head width

12.7(11.1-15.9)

Head height

15.4(14.2-16.6)

Snout length

6.3

6.3 (5.5-7.2)

Upper jaw length

13.6

13.2(12.5-13.9)

Diameter of pigmented eye

2.8

3.0 (2.6-3.4)

Diameter of pupil

1.8 (1.7-2.1)

Interorbital width

5.6

5.6 (5.1-6.0)

Posterior maxilla height

4.9

4.3 (3.2-5.4)

Postorbital length

18.2(17.6-18.6)

Preanal length

56.8

52.7 (48.8-56.8)

Predorsal length

28.4

28.1 (25.6-30.4)

Body depth at origin of anal fin

20.5

16.4(13.7-20.5)

Pectoral fin length

12.5

12.3(11.7-13.5)

Pectoral fin base height

6.4 (6.0-7.7)

Ventral fin length

22.7

23.3 (18.6-28.1)

Base ventral fin - anal fin origin

32.4(29.5-36.3)

* Data from Cohen and Hutchins (1982).

Description (Figs 35, 36).Tlie principal meristic and
morphometric characters arc shown in Table 15. Mature at
about 60 mm SL. Body slender, with pointed head profile.
Many cirri on snout. Eye size 2.6-3.4 (2.8)% SL. Head with
uniform and continuous squamation on cheeks, opercle and
occiput. Horizontal diameter of scales on body of 89 mm
SL female 1.3% SL, in about 33 horizontal rows. Maxillary
ending far behind eye, dorsal margin of maxillary covered
by upper lip dermal lobe, posterior end expanded. Anterior
nostril positioned high, 1/2.5 distance from upper lip to
anterior margin of eye. Posterior nostril moderately small,
about one-quarter the size of eye. Opercular spine with free
tip, pointed. Anterior gill arch with 19-26 (23) short rakers,
none elongate. Pseudobranchial filaments 2.

Head sensory pores (Fig. 36A-C). Supraorbital pores
7: 4 additional pairs of supraorbital pores as follows: 1
pair above and in front of eyes, a second above and behind
eyes, 2 more pairs on occiput in prolongation of upper
preopercular pore. Infraorbital pores 6 (3 anterior and 3
posterior), 3 posterior pores about half size of three anterior
pores. Mandibular pores 7-8 (3 anterior and 4-5 posterior):
first anterior pore large, tubular; one or two additional
posterior mandibular pores oriented in separate, though
incomplete, row parallel to original one and closer to jaw.
Preopercular pores: 3 lower, first and second with joined
opening, covered by dermal flap in lateral view, third non¬
tubular; upper preopercular pore present. [See description
of Alionematichthys ceylonensis for position of pores.]

Dentition (of holotype). Premaxilla with 11 outer
rows of granular teeth and 1 inner row of larger teeth.
Anteriormost teeth in inner row up to 1/5 diameter of
pupil. Vomer triangular, with 10 outer rows of small teeth
and 1 inner row of larger teeth up to 1/6 diameter of pupil.
Palatine with 7 outer rows of small teeth and 1 inner row
of larger teeth up to 1/6 diameter of pupil. Dentary with 9
outer rows of granular teeth and 1 inner row of larger teeth,
up to about 1/4 diameter of pupil.

Otolith (Fig. 36G-I). Moderately elongate in shape,
length to height 2.2 and moderately thick (otolith height to
otolith thickness about 2). Anterior tip rounded, posterior tip
rounded, slightly expanded. Dorsal rim thick, shallow, with
slight and broad depression at its middle. Inner face convex,
outer face slightly concave to flat, both smooth. Sulcus very
long, almost opening anteriorly, otolith length to sulcus
length 1.2. Sulcus slightly supramedially positioned and
slightly inclined, with separated colliculi and marked notch
at ventral rim of sulcus at joint of ostium with cauda. Ostium
length to cauda length 1.2-1.5. Ventral furrow distinct, close
to ventral rim of otolith.

Axial skeleton. Neural spine of vertebra 4 inclined and
5-10 depressed. Parapophyses present from vertebrae
6 to 14. Pleural ribs on vertebrae 2 to 13. First anal fin
pterygiophore much longer than subsequent ptcrygiophore,
reaching tip of last precaudal parapophysis.

Male copulatory organ (Fig. 36D-F). Two pairs of
pseudoclaspers, outer not extending beyond hood in resting
position. Outer pseudoelasper simple flap-shaped without
broadened base; inner pseudoelasper about length of outer
pseudoelasper or longer, reduced to stick-like inwardly
bent supporter. Isthmus moderately narrow; penis curved,
slightly longer than outer pseudoclaspers.

Colour. Live colour uniformly orange-brown (Allen
1985: 2268). Preserved colour uniformly light brown.

Comparison. See comparison between
Porocephalichthys and other genera above.

Distribution. Restricted in distribution to SW Australia,
around Perth and Rottnest Island (Fig. 30). The types were
collected in limestorc reef habitats, at depths of 3-15 m.

140

Dinematichthyine fishes of the Indo-west Pacific, Part IV

DISCUSSION

The fishes of the genera Dinematichthys and
Alionematichthys reviewed herein represent by far the
most common Dinematichthyini in the Indo-west Pacific,
comprising more than two-thirds of all investigated
specimens from the area. Both genera are particularly
common in reef core and outer reef environments, where they
often dominate the dinematichthyine samples, but are less
dominant in back reef / inshore and lagoonal environments.
They are entirely missing from locations outside of the
Indo-west Pacific reef belt (except for A. minyomma from
the Caribbean Sea). The tropical western Indian Ocean is
dominated by Dinematichthys iluocoeteoides ; less so in the
eastern Indian Ocean and western Pacific, where various
species of the genus Alionematichthys dominate. In the
latter region, the genus Diancistrus (see Schwarzhans
and Moller (2005)) usually occurs associated with, or as
competitor of, Alionematichthys and Dinematichthys. Other
dinematichthyine genera seem to be more adapted to reef-
associated niches in lagoon or near-shore environments (see
Schwarzhans and Moller (2007)).

Dinematichthys and Alionematichthys represent the
genera with the widest geographical distribution in the
tribe. Dinematichthys from the western Indian Ocean to
the western Pacific, absent only from the southeastern
Polynesian Islands; Alionematichthys is apparently not
found west of Sri Lanka, but reaches far east to Ducie Atoll,
Pitcairn Group, and constitutes the only dinematichthyine
genus which is represented in both the Indo-west Pacific
and the tropical western Atlantic by A. minyomma. Like
most other Dinematichthyini, the genera dealt with in this
last part of the review exhibit some restricted geographic
distribution patterns, but with fewer species than in the
similarly widely distributed genus Diancistrus. However,
the fishes dealt with in this review also contain some of
the most widespread species of the group, particularly
Dinematichthys iluocoeteoides (East Africa to Ogasawara)
and Alionematichthys piger (Andaman Islands to Ducie
Atoll). Alionematichthys riukiuensis is remarkable for its
large size. It can attain up to 150 mm SL, second in length
only to Dipulus caecus (up to 200 mm SL), but due to its
massive appearance it is certainly the foremost of all the
Dinematichthyini in body mass.

141

W. Schwarzhans and P. R. Muller

REGIONAL CHECKLIST

The purpose of the following regional check list is
to facilitate a quick approach for cross-checking when
analyzing a collection of dinematichthyinc fishes. The
regions are chosen to fit best with the geographic distribution
patterns observed in fishes of this group. Species considered
endemic to a particular region are underlined. It has to be
mentioned though that certain regions are under-represented
due to limited sampling, such as the Andaman Sea, the
Gulf of Oman, the Banda and Celebes Seas of Indonesia,
the Solomon Islands and Madagascar. It is recommended

Red Sea and Gulf of Aden

Dinematichthys ihiocoeteoides

Arabian Sea and Gulf of Oman

Dinematichthys ihiocoeteoides

East Africa south of 5°N

Dinematichthys ihiocoeteoides
Mascarenichthys sp.

Seychelles and Mascarenes

Dinematichthys ihiocoeteoides
Mascarenichthys heemstrai

Madagascar

Dinematichthys ihiocoeteoides
Maiuneaichthvs simplex

South Africa south of30°S

Dennatopsoides andersoni

Dermatopsoides kasougae

Dennatopsoides talboti
Mascarenichthys microphthalmus

Chagos Archipelago and Maldives

Diancistrus alleni
Dinematichthys ihiocoeteoides

Sri Lanka

Alionematichthvs cevlonensis

Dinematichthys ihiocoeteoides

Andamans, Thailand, Malaya and
Sumatra

Alionematichthvs phuketensis

A lionematichthys piger
A lionematichthys riukiuensis
Diancistrus sp.

Dinematichthys ihiocoeteoides
Eusiirciihis andamanensis

Ungiisurcuhis riauensis

Christmas and Cocos Islands

Dinematichthys trilohatus

that readers consult the check list of neighbouring areas
in these instances as well, since the lack of species is
probably artificial. Other areas have remained virtually
unsampled for Dinematichthyini, such as the tropical East
African shores, the Spratly Islands, Timor or the Natuna
Islands (South China Sea). Such areas could harbour further
unknown endemic species. Finally, we provide a key to all
currently recognised genera of Dinematichthyini. To key
out a species, readers need to use all six individual papers
in the revision: Moller et al. 2004a, Moller et at. 2005,
Schwarzhans et al. 2005, Moller and Schwarzhans 2006,
Schwarzhans and Moller 2007, plis the present paper.

Diancistrus macliidai
Diancistrus springeri
Dinematichthys ihiocoeteoides
Paradiancistrus cuvoensis

Unmtsurculus ohilippinensis

l /n (ms urea his williamsi

Celebes Sea

Alionematichthys piger
Alionematichthys riukiuensis
Diancistrus altidorsalis
Diancistrus heateae
Diancistrus karinae
Diancistrus macliidai
Dinematichthys ihiocoeteoides

Sunda Archipelago and
Indonesian Timor Sea

Beaglichthys bleekeri
Diancistrus alleni
Diancistrus altidorsalis
Diancistrus beateae
Diancistrus macliidai
Diancistrus novaeguineae
Diancistrus springeri
Dinematichthys ihiocoeteoides
Paradiancistrus lombokensis

Umriisurculus komodoensis

Ungusurculus sundaensis

Banda Sea and NW New Guinea

Alionematichthys piger
Alionematichthys plicatosurculus
Diancistrus alleni
Diancistrus altidorsalis
Diancistrus macliidai
Diancistrus nicer
Diancistrus novaeguineae
Dinematichthys ihiocoeteoides
Ogasawara Islands, Japan
Dinematichthys ihiocoeteoides

North Vietnam and Hainan
Island, China

Alionematichthys piger
Alionematichthys riukiuensis
Diancistrus vietnamensis

Kagoshima Islands, Japan

A lionematichthys piger
Alionematichthys riukiuensis
Diancistrus eiythraeus
Diancistrus fuscus
Dinematichthys ihiocoeteoides

Ryukyu Islands, Japan

Alionematichthys piger
Alionematichthys riukiuensis
A lionematichthys shinoharai

Diancistrus eiythraeus
Diancistrus fuscus
Diancistrus iackrandalli

Diancistrus sp.

Dinematichthys ihiocoeteoides

Taiwan and Philippines north of
15°N

Alionematichthys crassiceps
Alionematichthys piger
Alionematichthys riukiuensis
Brotulinella taiwanensis

Diancistrus eiythraeus
Diancistrus fuscus
Diancistrus macliidai
Dinematichthys ihiocoeteoides

Sulu Sea and Philippines south of

15°N

A lionematichthys piger
Alionematichthys plicatosurculus
Alionematichthvs suluensis

A lionematichthys riukiuensis
Diancistrus eiythraeus
Diancistrus fuscus
Diancistrus karinae

142

Dinematichthyine fishes of the Indo-west Pacific, Part IV

Belau, Guam, Caroline Islands
including Pohnpei

Alionematichthys crassiceps
Alionematichthys piger
Diancistrus atollorum
Diancistrus beciteae
Diancistrus karinae
Diancistrus pohnpeiensis
Dinematichthys iluocoeteoides
Marshall and Gilbert Islands
A lionematichthys piger
Diancistrus atollorum
Diancistrus beateae
Diancistrus mennei
Dinematichthys iluocoeteoides
Bismarck Archipelago and NE
New Guinea

Alionematichthys crassiceps
Alionematichthys piger
Alionematichthys plicatosurculus
Alionematichthys riukiuensis
Diancistrus alleni
Diancistrus altidorsalis
Diancistrus atollorum
Diancistrus beateae
Diancistrus eremitus
Diancistrus karinae
Diancistrus mcgroutheri
Diancistrus novaeguineae
Dinematichthys iluocoeteoides
Eusurculus pristinus
Ungusurculus collettei

Solomon Islands

A lionematichthys piger
Alionematichthys plicatosurculus
Alionematichthys riukiuensis
Diancistrus alleni
Diancistrus altidorsalis
Diancistrus beateae
Diancistrus eremitus
Diancistrus no vaeguineae
Dinematichthys iluocoeteoides
Eusurculus pristinus

NW Australia W of 142°E & N of
25°S

A lionematichthys piger
Alionematichthys riukiuensis
Alionematichthys sp.

Beaglichthys bleekeri
Beaglichthvs larsonae
Beaglichthys macrophthalmus
Brosmolus loneicaudus

Diancistrus alleni
Diancistrus beateae

Diancistrus jeffiohnsoni
Diancistrus novaeguineae
Didymothallus criniceps
Didvmothallus mizolepis
Dinematichthys iluocoeteoides
Dipulus hutchinsi
Eusurculus pistillum
Great Barrier Reef and SE New
Guinea

A lionematichthys piger
Alionematichthys riukiuensis
Alionematichthys sp.

Beaglichthys macrophthalmus
Diancistrus alleni
Diancistrus beateae
Diancistrus leisi
Diancistrus longifilis
Diancistrus mcgroutheri
Diancistrus novaeguineae
Didymothallus criniceps
Dinematichthys iluocoeteoides
Eusurculus pistillum
SW Australia VV of 145°E and S of
25°S

Dactvlosurculus somoni
Dermatopsoides morrisonae

Dipulus caecus
Dipulus hutchinsi
Dipulus multiradiatus
Porocephalichthvs dasvrhvnchus

Zephvrichthvs barrvi

SE Australia E of 145°E and S of
25°S

Dermatopsis hoesei
Dermatopsis macrodon

Monothrix polvlepis

Lord Howe Island

A lionematichthys piger
Diancistrus longifilis

New Zealand, North Island

Dermatopsis ioergennielseni

Norfolk Island

Dipulus norfolkanus

New Caledonia

A lionematichthys piger
A lionematichthys riukiuensis
Diancistrus longifilis
Didvmothallus pruvosti

Lapitaichthvs frickei

Loyalty Islands and Vanuatu

A lionematichthys crassiceps
Alionematichthys piger
Alionematichthys riukiuensis
Diancistrus alleni
Diancistrus beateae
Diancistrus brevirostris

Diancistrus longifilis
Diancistrus novaeguineae
Diancistrus tongaensis
Dinematichthys iluocoeteoides
Eusurculus pristinus
Paradiancistrus acutirostris

Fiji

Alionematichthys crassiceps
Alionematichthys piger
Alionematichthys winterbottomi
Dermatopsis vrecndeldi

Diancistrus beateae
Diancistrus eremitus
Diancistrus fiiiensis

Diancistrus rolmstus

Diancistrus tongaensis
Dinematichthys iluocoeteoides

Tonga

Alionematichthys crassiceps
A lionematichthys piger
A lionematichthys winterbottomi
Diancistrus alatus
Diancistrus mancioorus

Diancistrus tongaensis
Dinematichthys iluocoeteoides

Samoa

A lionematichthys samoaensis

Diancistrus alleni
Diancistrus beateae
Diancistrus tongaensis
Dinematichthys iluocoeteoides
Cook, Tubuai and Society Islands,
Pitcairn (inc. Ducie Atoll)
Alionematichthys piger
Diancistrus katrineae

Diancistrus tongaensis
Diancistrus sp.

143

W. Schwarzhans and P. R. Mollcr

Key to the genera of the Dinematichthyini

1 a. Head with continuous squamation on cheeks, opercle

and occiput.2

lb. Head with squamation patches on cheeks, and
occasionally also on opercle or without scales.3

2a. Precaudal vertebrae 14; dorsal fin rays >95; no canine
teeth; upper preopercular pore present; 3 pairs of pores
on occiput; sulcus of otolith with separate equally long

ostial and caudal colliculi ...

. Poroceplialiclithys

2b. Precaudal vertebrae 10-12; dorsal fin rays <95;
canine teeth present; upper preopercular pore absent;
no pores on occiput; sulcus of otolith with separate
colliculi, length ostial colliculum 2.5-4 times caudal
colliculum. Dinematichthys

3a. Anterior nostril positioned high (less than 1/3 the
distance from upper lip to aggregate distance to

anterior margin of eye). Alionematichthys

3b. Anterior nostril positioned low (1/3.5 to 1/6 the
distance from upper lip to aggregate distance to
anterior margin of eye).4

4a. Termination of maxilla low, not expanded.5

4b. Termination of maxilla expanded, angular or with
knob.8

5a. Opercular spine hidden.6

5b. Opercular spine exposed.7

6a. Single pair of pseudoclaspers with two equally long

supporters; upper preopercular pore present .

. Gunterichthys

6b. Single pair of pseudoclaspers with single supporter;
upper preopercular pore absent. Dermatopsoides

7a. Precaudal vertebrae 11-14; dorsal fin rays 64-85;

penis without hook near tip. Dermatopsis

7b. Precaudal vertebrae 13-25; dorsal fin rays 86-191;
penis with hook near tip. Dipulus

8a. A single pair of (outer) pseudoclaspers.9

8b. Two or three pairs of pseudoclaspers.13

9a. Two long supporters in pseudoclaspers.

Didymothallus

9b. Single supporter in pseudoclasper.10

10a. Precaudal vertebrae 11-12; sulcus of otolith with

separate colliculi .11

10b. Precaudal vertebrae 13-15; sulcus of otolith with
undivided colliculi . 12

1 la. Pseudoclasper simple flap with small hook at middle
of anterior rim. Lapitaichthys

lib. Pseudoclasper long and stick-like.

. Ogilbia mccoskeri

[note: only species of genus Ogilbia with single
pseudoclasper]

12a. Upper preopercular pore absent; dorsal fin rays
124-129; anal fin rays 90-94; total vertebrae 55-59;

D/A 37-42 . Brosmolus

12b. Upper preopercular pore present; dorsal fin rays
93-104; anal fin rays 64-76; total vertebrae 45-47;
D/A 28-35. Monothrix

13a. 3 pairs of pseudoclaspers.14

13b. 2 pairs of pseudoclaspers. 15

14a. Inner pseudoclasper separated into an anterior and a
posterior pseudoclasper, pseudoclaspers not joined at
base; 6-7 branchiostegal rays; 11 precaudal vertebrae;
pectoral fin rays 16-21; first and second lower

preopercular pore joined in a single opening.

. Ogilbichthys

14b. Second inner pseudoclasper inserted between
first inner pseudoclasper and outer pseudoclasper;
pseudoclaspers joined at base; 8-9 branchiostegal
rays; 13-14 precaudal vertebrae; pectoral fin rays
22-26; first and second preopercular pores with

separate openings. Dactylosurcidus

15a. Upper preopercular pore absent.16

15b. Upper preopercular pore present .20

16a. No scales on head; specimens longer than 20 mm
SL without visible eyes, only minute black dots in
specimens less than 20 mm SL. Typhliasina

16b. Scales on cheeks; all specimens with visible eyes ...

.17

17a. First and second preopercular pores with separate
openings.18

17b. First and second lower preopercular pore joined in a
single opening.19

18a. Inner pseudoclasper anteriorly connected to outer
pseudoclasper; sulcus of otolith with undivided
colliculi. Beagliclithys

18b. Inner pseudoclasper branched, medially connected to
outer pseudoclasper; sulcus of otolith with separate
colliculi, cauda short. Zephyrichthys

19a. Inner pseudoclasper concave, anteriorly connected to

outer pseudoclasper; precaudal vertebrae 11.

. Diancistrus manciporus

[note: only species of genus Diancistnis without upper
preopercular pore]

19b. Inner pseudoclasper claw-like; precaudal vertebrae
12. Ungusurculus

144

Dinematichthyine fishes of the Indo-west Pacific, Part IV

20a. Single lower preopercular pore.21

20b. Three lower preopercular pores, but first and second
pore open into single opening.22

21a. Inner pseudoclasper about as large as outer

pseudoclasper and separate. Pseudogilbia

21b. Inner pseudoclasper half the size ofouter pseudoclasper

and anteriorly connected to U-shaped structure.

. Paradiancistms

22b. Inner pseudoclasper anteriorly joined to outer

pseudoclasper.23

22a. Inner and outer pseudoclaspers free.24

23a. Body slender (head height < 15% SL. depth at anal

< 16% SL); precaudal vertebrae predominantly 12
(rarely 11); maxilla rounded posterior-ventrally with
weak knob in front of rear comer; body scales small,

< 1.2% SL. Brotulinella

23b. Body robust, moderately slender to deep (head

height > 15% SL except for Diancistrusjeffjohnsoni,
body depth at anal > 16% SL except for Diancistrus
longijilis ); precaudal vertebrae 11 (except 12 in
Diancistrus jeffjohnsoni); maxilla with angular
postero-ventral widening close to its termination;
body scales (1.2) 1.3—2.2% SL . Diancistrus

24a. Inner pseudoclasper hidden in pocket of isthmus when

in resting position; penis hooked.

. Mascarenichthys

24b. Inner pseudoclasper open; penis bent, but not
hooked.25

25a. Inner pseudoclasper sucker-disk like; supporter of
outer pseudoclasper distally expanded and usually

with anterior hook. Eusurculus

25b. Inner pseudoclasper simple flap or diversified, but not
sucker-disk like; supporter of outer pseudoclasper not
expanded and without anterior hook.26

26a. Sulcus of otolith with separated colliculi ....Ogilbia

26b. Sulcus of otolith with undivided colliculi.

. Majungaichthys

ACKNOWLEDGMENTS

We wish to thank the following people for helping us
with material and information: Gerald R. Allen (WAM),
M. Eric Anderson (SAIAB), Dianne J. Bray (NMV),
David Catania (CAS), Daniel M. Cohen (CAS), Gavin
Dally (NTM), Guy Duhamcl (MNliN), Jon Fong (CAS),
Kiyoshi Hagiwara (YCM), Karsten Hartel (MCZ), Philip C.
Hecmstra (SAIAB), Erling Holm (ROM), Peter A. Hulley

(SAM), J. Barry Hutchins (WAM), Tomio Iwamoto (CAS),
Susan L. Jewett (USNM), Jeffrey W. Johnson (QM), Helen
K. Larson (NTM), Jeff M. Leis (AMS), Yoshihiko Machida
(BSKU), James Maclaine (NHM former BMNH), Mizuki
Matsunuma (KAUM), John E. McCosker (CAS), Mark
A. McGrouther (AMS), Sue Morrison (WAM), Hiroyuki
Motomura (KAUM), Vusi Mthombeni (SAIAB), Jorgen G.
Nielsen (ZMUC), John R. Paxton (AMS), Patrice Pruvost
(MNHN), John E. Randall (BPBM), Sandra Raredon
(USNM), Sally Reader (AMS), Clive Roberts (NMNZ),
Robert H. Robins (UF), Mark Sabaj (ANSP), Jeff Seigel
(LACM), Gento Shinohara (NSMT), Shirleen Smith
(USNM), David G. Smith (USNM), William F. Smith-Vaniz
(USGS), Victor G. Springer (USNM), Andrew L. Stewart
(NMNZ), Arnold Suzumoto (BPBM), Tom Tmski (AMS),
Jeffrey T. Williams (USNM) and Richard Winterbottom
(ROM). Special thanks go to Ronald Fricke (SMNS) and
Helen K. Larson (NTM), for careful reviews of all four
papers on the Dinematichthyini of the Indo-west Pacific.

Also thanks are due to the following colleagues at
ZMUC: Brigitte Rubaek for making the specimen drawings,
Geert Brovad for producing the photos, and Tammes
Menne for help with x-raying and packing. The project was
financed by the Carlsberg Foundation and by the Visiting
Collection Fellowship grant from the Australian Museum,
Sydney.

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Accepted 6 November 2008

146

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 147-150

A new species of the gudgeon Bostrychus (Teleostei: Gobioidei: Eleotridae),

from peninsular Malaysia

HELEN K. LARSON

Museum and Art Gallery Northern Territory,

Department of Natural Resources, Environment, the Arts and Sport,

GPO Box 4646, Darwin, NT0801, AUSTRALIA
helen.larson@nt.gov.au

ABSTRACT

A new species of the elcotrid genus Bostrychus is described from a single specimen obtained from a disturbed mangrove
site at the Sementa River, in Selangor State, Malaysia. It resembles B. sinensis and B. strigogenys in vertebral number,
body shape and interorbital pore pattern, but differs from all known species of Bostrychus by the ladder-like barred
body markings, the spotted, dark-margined pectoral fins and low fin ray counts. It is the second known “estuarine”
species of the genus.

Keywords: Eleotridae, Bostrychus, new species, estuarine, Malaysia.

INTRODUCTION

There are six known species of the eleotrid genus
Bostrychus Lacepede: five from the western Pacific and one
from west Africa, with the last species sometimes placed
in the genus Hannoichthys Herre. Hoese and Kottelat
(2005) discussed these species in their description of the
cave-dwelling B. microphtlialmus and provided a key to the
Pacific species. Bostrychus greatly resembles Odonteleotris
Gill, but for the possession of vomerine teeth (lacking in
Odonteleotris) and difference in adult size ( Odonteleotris
reaching at least 305 mm SL, Bostrychus reaching 185 mm
SL) (pers. obs.; Hoese and Kottelat 2005).

No species of Bostrychus is very well studied, but three,
B. africanus, B. sinensus and B. zonatus, arc regularly
recorded from estuarine (mangrove) habitats. The other
species all appear to be confined to fresh waters and are
highly localised in distribution. This paper describes a new
species from a mangrove habitat in Malaysia. The collection
site was searched for additional specimens subsequent to
the new species’ discovery, but none were found. The site
is now ‘reclaimed’ land.

METHODS

Measurements were taken using electronic callipers
and dissecting microscope. Counts and methods generally
follow Hubbs and Lagler (1970), except as indicated below.
Transverse scale counts backward (TRB) arc taken by
counting the number of scale rows from the anal fin origin
diagonally upward and back toward the second dorsal fin

base. Head length is taken to the upper attachment of the
opercular membrane. The segmented or branched caudal
ray pattern (e.g. 9/8 or 9/7) is the number of segmented
caudal rays attaching to the upper and lower hypural
plates respectively. Vertebral counts and other osteological
information was obtained by X-ray. Pterygiophore formula
follows Birdsong et al. (1988).

Abbreviations. NTM - Museum and Art Gallery of
the Northern Territory (previously Northern Territory
Museum), Darwin.

Comparative material. Bostrychus sinensis, NTM
S. 12731 -018,4(44-90), Yonada River, Iriomote-jima, Japan;
NTM S. 10555-001, 2(79-92), Ludmilla Creek, Darwin,
Northern Territory, Australia. Bostrychus zonatus, NTM
S.14219-001, 1(86), Ludmilla Creek, Darwin, Northern
Territory, Australia; NTM S. 14768-001, 2(93-114), creek
olTDick Ward Drive, Ludmilla, Darwin, Northern Territory,
Australia.

SYSTEMATICS

Bostrychus Lacepede, 1801

Bostrychus Lacepede, 1801: 140 (type species
Bostrychus sinensis Lacepede, 1801, by subsequent
designation).

Boroda Herre, 1927: 58 (type species Boroda expatria
Herre, 1927, by original designation).

Hannoichthys Herre, 1950: 198 (type speces Eleotris
africana Steindachner; replacement for llatino Herre, 1946,
preoccupied by Hanno Gray, in Mammalia).

H. K. Larson

Bostrychus scalaris sp. nov.

Ladder gudgeon

Material examined. HOLOTYPE - NTM S. 15552-003,
93 mm SL male, small pool at low tide, disturbed mangrove
at Sementa River, 3° 4.84’ N 101° 21.35’ E, near Klang,
Selangor State, Malaysia, H. Larson, A. Sasekumar, S. Lim,
G. Liew and parasitology students, 5 October 2002.

Diagnosis. A Bostrychus with 1,9 second dorsal fin rays;
1,8 anal fin rays; 16 pectoral rays and 135 to 142 lateral
scales; mandibular sensory' papilla row / arranged in clusters;
broad, depressed head and slender, compressed body;
heavily pigmented pectoral fins, and distinctive ladder-like
dark bars and complex serpentine pattern on the body.

Description. Based on the male holotype, 93 mm SL
(Figs 1, 3).

First dorsal spines VI; second dorsal rays 1,9; anal rays
1,8; pectoral rays 16 (on both sides); segmented caudal
rays 17, in 9/8 pattern; branched caudal rays 8/7; lateral
scale count 135 (on left side; 142 on right); transverse
scales backward 42 (on left; 35 on right); predorsal scale
count 32; vertebrae 12+15; dorsal pterygiophore pattern
3-3210; 2 epurals; 4 anal pterygiophores anterior to first
haemal spine.

Body slender, somewhat rounded anteriorly, compressed
posteriorly; body depth at anal fin origin 15.5% of SL.
Caudal peduncle long, length 23.0% of SL. Caudal peduncle
depth 12.3% of SL. Head broad, depressed, considerably
wider than deep at preopcrcular margin, head length 25.7%
of SL; head depth at posterior preopereular margin 46.4%
of HL; width at posterior preopereular margin 75.7%
of HL; greatest width, at inflated cheeks, 82.4% of HL.
Mouth large, terminal and oblique, forming an angle of
about 30° with body axis; jaws ending at point just behind
eye. Upper jaw length 56.5% of HL; inner margin of lips
finely fimbriate; lower lip fused to chin anteriorly, side of
lip free; chin slightly inflated anterior to groove containing
mental papillae. Anterior naris at end of long tapering tube at
edge of upper lip; posterior naris oval, in short tube sunken
in distinct pit; adjacent to anterdorsal margin of eye. Eye
small, lateral, width 15.9% of HL. Interorbital broad, fleshy,
37.7% of HL. Snout broad, flat, blunt anteriorly in dorsal
view, flattened, slightly pointed in lateral view, 32.2% of
HL. Gill opening damaged (removed for monogenean
parasite study) but judging from tissue remnants, opening
extending forward to under posterior margin of preopercle.
Tongue large, tip gently rounded. Teeth in both jaws small,
evenly sized, conical and rather blunt-tipped; in six or

Fig. I . Living captive holotype of Bostrychus scalaris, NTM S. 15552-
003, 93 mm SL male, Sementa River, Malaysia, showing distinctive
barred colour pattern. Image has been digitally modified, to remove
white reflective scratches (on plastic aquarium wall).

148

seven rows across front of jaws, and four or five rows
along side. Vomerine teeth low, blunt, conical, arranged in
broad triangular patch six to eight rows deep. Headpores
as shown in Figures 4, 5; five preopcrcular pores present;
single median anterior interorbital pore; paired posterior
interorbital pores; anterior (small and close to edge of upper
lip) and posterior nasal pores; six pores from behind eye in
broken oculoscapular canal. Sensory papillae in transverse
pattern (Figs 4,5); mandibular papillae i (sensu Sanzo 1911)
in few short transverse rows and mostly arranged in eight or
nine irregular clusters; pair of / papillae clusters on chin.

Body covered with small cycloid scales, reaching onto
head; scales on head largely embedded in fleshy skin, can
be difficult to discern. Opercle covered with small cycloid
scales. Preopercle with small cycloid scales covering most
of check (difficult to see. especially near papilla rows).
Prepelvic region with embedded small cycloid scales
extending forward onto isthmus (actual extent indeterminate
due to damage). Pectoral fin base covered with small cycloid
scales. Predorsal scales small, cycloid, extending up to just
above opercle. Belly with embedded cycloid scales.

First dorsal fin low, pointed (tips of third and fourth
spines twisted), adpressed fin falling well short of second
dorsal fin spine, with gap of about 17 small scales between
the two fins. Postcriormost second dorsal and anal rays taller
than first dorsal fin but not greatly so; posterior rays slightly
longer than anterior rays but not greatly so, posterior rays
falling well short of caudal fin base. Pectoral fin broadly
rounded, central rays longest, 16.9% of SL; upper and
lowermost two rays unbranched. Pelvic fin length 18.0%
of SL; pelvic fins slender, pointed, fourth rays longest, fins

Fig. 2. Living captive holotype of Bostiychus scalaris, close-up of
head.

Fig. 3. Preserved Bostiychus scalaris holotype, NTM S. 15552-003
(attitude due to gill arches being removed for parasite study).

A new species of the gudgeon Bostrychus

extending less than half the distance to anus. Caudal fin oval,
rounded posteriorly; caudal fin length 22.7% of SL.

Live colouration. Based on photographs of captive
holotypc (Figs 1,2), taken under field conditions (not ideal;
and using 2001-vintage digital camera). Mead and body
whitish with dark brown and pale gold markings. Head
pinkish white on lower half, with dark brown to pale gold,
ocellate to vermiculate, convoluted pattern covering most
of head, markings coalescing dorsally; darkest and most
conspicuous part of pattern a blackish brown broad band
from snout running along suborbital and joining similar
blackish brown bar from eye. Iris red-gold, outer margin
of iris brown-speckled, thin light golden edge around pupil.
Mid-side of body with ladder-like pattern of 24 short vertical
dark brown bars, all joined together ventrally by irregular
dark brown, partly broken, broad line and coalescing
dorsally with serpentine pattern of broad dark brown lines
and spots on dorsum; bars and spots and vermiculate lines
mostly outlined diffusely with pale gold. Upper part of
caudal fin with round black spot broadly ocellated with
dark yellow; remainder of fin dark yellow with about five
irregular bands of dark brown, outermost row blackish and
reaching posterior margin of fin. First dorsal fin mostly
folded down in only available photographs; blotches of
dark brown and dark yellow partly visible. Second dorsal
fin yellow with about seven oblique irregular bands of
dark brown to blackish brown. Anal fin pale yellowish
white; some purplish brown and dull yellowish mottling
may be present (unclear from photos). Pectoral fin reddish
orange with broad dark brown bar curving along fin base
and outer half of fin blackish brown with narrow whitish
margin, remainder of fin with dark brown spots and streaks
following fin rays. Pelvic fins whitish with broad dusky
streak or blotch on fin rays (not clear in photos).

Colouration in alcohol. Pattern same as live colour
but body whitish with greyish brown markings on head
and body (Fig. 3). Distinct slightly oval blackish spot,
surrounded by white, on upper caudal fin base. Caudal
fin spots dark brown. Pectoral fin markings blackish, with
broad fin margin darkest. Anal fin white with indistinct
scattered fine grey elongate marks running along fin rays.
Pelvic fins whitish with broad dusky grey streak along
central rays.

Fig. 4. Lateral view of head of Bostrychus scalaris holotype, showing
lateral canal pores and sensory papillae.

Fig. 5. Bostiyclius scalaris holotype. A, dorsal view of head of
showing lateral canal pores; B. ventral view of part of head, showing
mandibular sensory papillae (gill region damaged).

Distribution. Known from only one specimen from
Selangor state on the west coast of peninsular Malaysia,
and from a watercolour painting (Fig. 6) in a notebook
compiled in Singapore between 1858 and 1862 by the
French naturalist F. L. de Castelnau, and now stored in
the Zoological Museum of the University of Liege in
Belgium (Loneux 2006). No specimens from Singapore
are known.

Comparisons. This new species differs considerably in
colour pattern from all other known species of the genus,
having strongly pigmented pectoral fins, unlike all the
others, which all have relatively plain unspotted pectoral
fins, and it has a distinctive barring and serpentine pattern
on the body. The ladder-like pattern of 24 short vertical
dark bars along the side of the body somewhat resembles
the bars seen in B. zonatus Weber, but this species has
seven to nine broad brown bars along the lower half of the
body and finely speckled and spotted dorsum. Good colour
photographs showing the body pattern of living specimens
of B. strigogenys Nichols and B. zonatus can be seen in
Allen et al. (2000).

Bostrychus scalaris, B. sinensis Lacepede and
B. strigogenys are all similar in body shape, in that they
have broad flat heads with long jaws, while B. zonatus has
a short snout and jaws and more cylindrical head; they also
share similar first dorsal fin spine counts (VI-V1II) and
vertebral numbers (26-30), while B. zonatus has V1II-X first
dorsal fin spines and 36-38 vertebrae (pers. obs; Hoese and
Kottelat 2005). The sunken pit in which the posterior naris
protrudes from in B. scalaris is also present in B. sinensis
and B. zonatus (other species not examined).

The mandibular sensory i papillae are arranged in
proliferated clusters, with only the posteriormost three rows
arranged in what could be described as short transverse
rows, and the row is mostly set in a deep fleshy groove.
This pattern is similar to that observed in B. sinensis and B.
zonatus in which the transverse / rows are partly doubled or
proliferated, but not to such an extent (pers. obs.).

Ecology. The holotype was found under a small log
in a very small muddy pool at low tide, in a partly cleared
mangrove (dominated by Avicennia marina) on the Semcnta

149

H. K. Larson

Fig. 6. Castelnau’s painting of Bostrychus scalaris. not identified to genus, from his Singapore notebooks.
Photograph courtesy University of Liege, Belgium.

River, near Klang. The adjoining area was searched twice
for additional specimens. The collection site is now
“reclaimed” land. The water was brackish (20 ppt) and only
a few centimetres deep.

The gill arches were removed for an ongoing survey of
monogenean parasites and their relationships; unfortunately
none were found (S. Urn, pers. comm.).

Etymology. From the Latin scalaris , a ladder, in
reference to the step-ladder-like banded pattern on the
body of this species. A suggested common name is ladder
gudgeon.

Remarks. This species must have existed in Singapore
at one time, as it was collected and illustrated by Castelnau
(Fig. 6), some time between 1858-1862. This fish was
identified only by the number 607, with no notes as to what
genus Castelnau thought it might be. It is not known what
has become of the specimen. The only Bostrychus that has
been reported from Singapore is B. sinensis (Larson el al.
2008).

ACKNOWLEDGMENTS

My many thanks to Professors Susan Lim and
A. Sasekumar of the University of Malaya, and to their
students, for all their help in finding this fish and for
their companionship. And thanks to Barry Russell of
Marine Biodiversity (Department of Natural Resources,
Environment, the Arts and Sport, Northern Territory
Government), for bringing my attention to Castelnau’s long-
lost paintings at the University of Liege, Belgium.

REFERENCES

Allen, G.R.. Hortle, K.G. and Renyaan, S.J. 2000. Freshwater fishes
of the Timika region New Guinea. P.T. Freeport Indonesia,
Timika.

Birdsong, R.S., Murdy, E.O. and Pezold, F.L. 1988. A study of the
vertebral column and median fin osteology in gobioid fishes
with comments on gobioid relationships. Bulletin of Marine
Science 42(2): 174—214.

Herre, A.W.C.T. 1927. Gobies of the Philippines and the China
Sea. Monographs of the Philippine Bureau of Science,
Manila.Philippine Islands. Monograph 23: I 352.

Herre, A.W.C.T. 1946. New genera of Eleotridae and Gobiidae
and one new species from West Africa. Proceedings of the
Biological Society of Washington 59: 121-126.

Herre, A.W.C.T. 1950. A new name for Hanno. a genus of African
gobies. Stanford Ichthyological Bulletin 3(4): 198.

Hoesc, D.F. and Kotlelat, M. 2005. Bostrychus microphthalmus, a
new microphthalmic cavefish from Sulawesi. Ichthyological
Exploration ofFreshwaters 16(2): 183-191.

Hubbs, C.L. and Laglcr, K.F. 1970. Fishes of the Great Lakes Region.
University of Michigan Press: Ann Arbor.

Lacepede, B.G.E. 1801. Histoire naturelle des poissons. Volume 3:
i-lxvi, 1-558. Paris.

Larson. H.K., Jafaar, Z. and Lim, K.P. 2008. An annotated checklist
of the gobioid fishes of Singapore. Raffles Bulletin of Zoology
56(1): 135-155.

Loneux. M. 2006. The Castelnau's fish collection and watercolour
notebooks. Pp. 91-94. In Scgers, H., Desmct, P. and Baus,
E. (eds) Tropica! biodiversity: science, data, conservation.
Proceedings of the 3rd GB1F Science Symposium, Brussels,
18-19 April 2005, 169 pp.

Sanzo, L. 1911. Distribuzionedellepapillecutanee(organi ciatifonni)
e suo valore sistematico nei Gobi. Mitteilungen aus der
Zoologischen Station zu Neapel 20: 249-328.

Accepted 23 July 2008

150

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 200S 24: 151-155

Diversity of bats at two contrasting elevations in a protected dipterocarp forest in

Sarawak, Borneo

J. MOHD-AZLAN 1 - 2 , SITI HASMAH TAHA 2 , CHARLIE J.M. LAMAN 2 AND M.T. ABDULLAH 2

1 School for Environmental Research, Charles Darwin University, Casuarina, NT 0909, AUSTRALIA
2 Department of Zoology), Facility of Resource Science and Technology, Universiti Malaysia Sarawak,

43000 Kota Samarahan, Sarawak, MALAYSIA
Corresponding author: azlan@cdu.edu.au

ABSTRACT

We present an assessment of the diversity of Bornean bats at two contrasting elevations (119 m and 787 m) in Kubah
National Park surveyed between August and December 2006. Three hundred and eighty-two individuals of bats from
26 species representing six families were recorded using 20 mist nets and three harp traps. The most commonly caught
bat was Hipposidems cervinus (Gould, 1863) (n= 168) followed by Penthetor lucasi (Dobson, 1880) (n = 55), and they
were also the most commonly detected species at low and high elevation sites, respectively. This survey yielded the first
recorded specimen of Pipistrellus cuprosus Hill and Francis, 1984 for Sarawak. Analysis of daily cumulative capture rates
indicated that further effort at these sites might not yield additional species if sampling techniques and locations were
maintained. The species diversity index of Megachiroptcra was higher at high elevations (H 0.567 versus H’ = 0.466),
whereas the species diversity index of Microchiroptera was greatest at low elevations (H'= 0.905 vs IT = 1.225).

Keywords: Borneo, Chiroptera, bat diversity, elevation.

INTRODUCTION

Bats play a major role in ecosystem processes including
pollination, seed dispersal and forest insect regulation
(Davidson and Zubaid 1992; Tan et al. 1998; Payne
et al. 2005). The importance of bats in these processes
is particularly significant in tropical rainforests where
chiropteran richness and diversity is high. Malaysia is
renowned for these attributes of its bat fauna, however they
vary with location and forest type. For example, Francis
(1990) recorded 44 chiropteran species including nine
species of fruit bats and 35 microbats in a primary lowland
dipterocarp forest at Pasoh Forest Reserve, peninsular
Malaysia, while Hall et al. (2002) recorded 23 species of
bats (six Megachiroptcra, 17 Microchiroptera) in a primary
mixed dipterocarp forest (lowland mixed dipterocarp forest
and tall mixed dipterocaip forest) at Niah National Park,
Sarawak, Borneo. Despite tropical rainforests supporting
high levels of bat diversity (Corbett and Hill 1992; Payne
et al. 2005), studies on Bornean diversity arc lacking
(Struebig et al. 2006) although such basic information is
important and plays a vital role for species conservation
management, especially of rare and cryptic species
(Frey 2006). Regional degeneration, fragmentation and
deforestation pose a major threat to bats in South-east Asia
(Lane et al. 2006). In light of this, we report the findings of
our survey of Kubah National Park in Sarawak, Borneo.

METHODS

Study area. Kubah National Park (Fig. 1) is situated
20 km from Kuching city and covers an area of 2742 ha.
This protected area consists of mixed dipterocarp forest,
riverine forest, montane forest and heath forest. Several
longhouses around this National Park have caused intrusion
into, and excision of, forest produce. The terrain throughout
this Park is steep with high ridges. The emergent tree
species in the Park arc forest species of the genera Shorea,
Dryobalanops and Dipterocarpus. Other dominant tree
species include Calophyllum sp„ Cotylelohium malayanum
v. Slooten and Litsea resinosa Blume. Details on the
vegetation composition have been described by Hazebroek
and Abang Morshidi (2000). Bcnnet and Walsh (1988) only
recorded 18 terrestrial mammal species, suggesting this
Park has relatively low large mammal diversity compared
to other protected areas in Sarawak (Mohd-Azlan et al.
2007). Sampling was concentrated at two elevations, 119
m and 787 m above sea level, for 14 non-consecutive days
between August and December 2006. The lower elevation
site consisted mostly of lowland dipterocarp forest with
a mixture of primary and old secondary forest, while
the higher elevation site was a mixture of primary hill
dipterocarp forest and lower montane forest.

Sampling Methods. Each night, twenty mist nets and
three two band-harp traps were set up at each elevation for

J. Mohd-Azlan, S. H. Taha, C. J. M. Laman and M. T. Abdullah

Fig. 1. Montane forest in Kubah National Park provides habitat for
bats. Photograph by J. Mohd-Azlan.

12 hours, from 1830 hr until 0630 hr. Bats were captured
by using 36 mm mesh mist nets measuring 9 m and 12 m
in length. These nets were erected in the forest understorey
(up to 1 m above the ground), across major trails, rivers
and clearings in the forest and at the forest edge. Nets and
traps were relocated to new sites every two to four days.
Identification was based on Payne el al. (2005). Bats caught
were identified and released immediately. To examine and
compare species diversity, the H' values of the Shannon-
Weiner Index (ranging from 1 for high species diversity
to 0 for low species diversity) were calculated for both
Megachiroptera and Microchiroptera at lower and higher
elevation We also used Zars T-test (Zars 1996) to test the
difference in species abundance between high and low
elevation.

RESULTS AND DISCUSSION

Twenty-six species, representing 28.0% of the 92
chiropteran species recorded in Borneo (Payne el al. 2005),
were captured during our survey (Table 1). This included
eight species of Megachiroptera (47.1 % of Bornean species)
and 18 species of Microchiroptera (24.0% of Bornean
species) representing six families. The most commonly
encountered bat at lower elevation was Hipposideros
cervinus (Gould, 1863) (n = 168, or 63.8% of total capture),
while Penthetor lucasi (Dobson, 1880) dominated captures
at higher elevation (n= 55, or 45.1 % of total capture). Most
of the adult female P. lucasi captured were either pregnant
or lactating. Significantly, our survey also obtained the first
recorded capture of Pipistrellus cuprosus Hill and Francis,
1984 for Sarawak (Fig. 7).

The total number of Megachiroptera caught at the lower
elevation was 57 (H ’= 0.466), while at the higher elevation
the total number captured was 107 (H’= 0.567). Whereas
the total number of captures of Microchiroptera at the lower
elevation was 203 (H ’= 1.225), we only recorded 15 (//’
= 0.905) at the higher elevation. The Zar’s (1996) t-test on
H' value of Shannon-Weiner Index had been calculated
for both Megachiroptera and Microchiroptera at both the
lower and the higher elevations, resulting in a 0025 ,299=

2.44 (t = -2.4409, df= 300, p<0.025) and showing that there
are significantly less species present at higher elevations
compared to lower elevations. However, due to the short
sampling periods, and without involving replicates, our
comparative results are limited, but they do provide baseline
data for future studies.

In general, highland areas in tropical rainforests support
relatively low chiropteran species diversity compared to
lowland forests (Tuen el al. 2002). For example, Salleh
etal. (1999) recorded only five species of bats from Kelabit
Highlands, whereas Mohd-Azlan et al. (2003) recorded 11
species at Kayan Menterang National Park, East Kalimantan.
In addition to this, the species accumulation curve (Fig. 2)
reaches an asymptote after 10 nights suggesting that almost
all of the understorey chiropteran species subject to these
trapping methods in this area were recorded. However, the
overall species diversity may not represent the bat fauna of
the entire Kubah National Park due to the limited number
of sampling methods, duration of the study, and types and
structure of forest. These factors directly affect the number
of species and individuals that are likely to be captured
(Kingston et al. 2003). Further surveys targeting open
spaces, forest edges and the canopy are likely to record
species of other foraging guilds (Struebig et al. 2006). In
view of this, rapid assessment of the bat fauna (especially
the Microchiroptera) using conventional capture methods
alone may not provide comprehensive information on the
chiropteran diversity in tropical rainforests, thus under¬
representation of microchiropteran faunal diversity can be
expected in the majority of environmental assessments in
Malaysia.

Fig. 2. Species accumulation curve of species richness against
sampling night in Kubah National Park, Sarawak.

ACKNOWLEDGMENTS

We would like to thank Universiti Malaysia Sarawak
for financial support, Sarawak Forestry Department and
Sarawak Forest Corporation for their assistance in allowing
access into Kubah National Park and permits (No. 41 /2006).
We are also grateful to Mr Mohidin Rajuli, Mr Jack Derring,

152

Bat diversity in tropical rainforest Borneo

Table 1. Number of individuals and relative abundance of bats captured at both elevations at Kubah National Park between August and
December 2006.

No. Species

Loner (n)

Relative
abundance (%)

Upper (n)

Relative
abundance (%)

Family Pteropoiiidae

1. Cynopterus brachyotis { Muller, 1838)

13.68

17.22

2. Megaerops ecaudatus (Temnick. 1837) (Fig. 3)

0.38

16.40

3. Balionycteris maculata ( Thomas, 1893)

1.52

4.10

4. Penthetor lucasi ( Dobson. 1880)

4.94

45.10

5. Eonycteris spelaea (Dobson. 1871)

0.76

6. Chironax melanocephalus (Temnick, 1825) (Fig. 5)

0.38

7. Macroglossus minimus (E. Geoffroy, 1810)

0.82

8. Cynopterus horsefieldi Gray, 1843

4.10

Family Emballonuridae

9. Emballonura alecto (Eydoux and Gervais, 1836)

0.38

0.82

10. Embalionura monticola Temnick, 1838

0.82

Family Nvcteridae

11 . Nycteris tragata (K. Andersen, 1912)

0.38

Family Rliinolophidae (Fig. 4)

12. Rhinolophus borneensis Peters, 1861

0.38

13. Rhinolophus arcuatus Peters, 1871

0.82

14. Rhinolophus affinis Horsfield, 1823

0.76

2.46

15. Rhinolophus sedulus K. Andersen, 1905

0.76

1.64

Family Hipposideridac

1 6. Hipposideros ater Templeton, 1848

3.80

0.82

17. Hipposideros bicolor (Temnick, 1834)

0.38

18. Hipposideros dyacorum (Temnick, 1834)

0.38

19. Hipposideros ridleyi Robinson and Kloss, 1911

2.28

20. Hipposideros cervinus (Gould, 1863)

168

63.84

0.82

21. Hipposideros galeritus Cantor, 1846

0.38

1.64

Family Vesperlilionidae

22. Myotis ridleyi Thomas, 1898

0.38

23. Pipistrellus cuprosus Hill and Francis, 1984 (Fig. 7)

2.46

24. Kerivoulapapillosa (Temnick, 1840) (Fig. 6)

1.14

25. Kerivoula hardwickii (Horsfield, 1824)

1.52

26. Marina suilla (Temnick, 1840)

0.38

Total records

260

100

122

100

Number of species

Number of families

Net / trap-hour

6624

Bats/ effort (Net/trap hour)

0.039

0.018

S-\V Index (Mcgachiroptcra)

0.467

0.567

S-W Index (Microchiroptera)

0.369

0.905

Simpson (1-D) (Mcgachiroptcra)

0.848

0.841

Simpson (1-D) (Microchiroptera)

0.938

0.943

Evenness (Mcgachiroptcra)

0.528

0.399

Evenness (Microchiroptera)

0.733

0.081

153

J. Mohd-Azlan, S. H. Taha, C. J. M. Laman and M. T. Abdullah

Figs 3-7. Representatives of bats captured in Kubah National Park: 3, Megaerops ecaudatits; 4, Rhinolophus sp; 5, Chironax melanocephalus;
6, Kerivoula papillosa; 7, Pipislrelltis cuprosus. Photos: 3, 4, J. Mohd-Azlan; 5, 6, 7, S. H. Taha.

154

Bat diversity in tropical rainforest Borneo

Dr Andrew Alek Tuen, Ms Suziani Sulaiman, Mr Wahap
Marni, Mr Besar Ketol, Mr Jailani Mortada, Mr Huzal
Irwan Husin and Ms Ratnawati Hazali for the assistance
in the field. Comments from Dr Damian Milne are greatly
appreciated. We also thank the three anonymous reviewers
for their helpful comments on a draft of this manuscript.

REFERENCES

Bennet, E.L. and Walsh, M. 198B. A wildlife survey of the proposed
Matang National Park, Sarvvak. Unpublished report. National
Parks and Wildlife Office, Forest Department, Kuching
Sarawak.

Corbet. G.B. and I fill, J.E. 1992. The mammals of the Indomalayan
Region: a systematic review. Oxford University Press: New
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Davidson, G.W.ll. and Zubaid, A. 1992. Food habits of the lesser
false vampire bat. Megaderma spasma from Kuala Lompat,
peninsular Malaysia. Zeitschrift fur Saugetierkunde 57:
310-312.

Francis, C.M. 1990. Trophic structure of bat communities in the
understorey of lowland dipterocarp rain forest in Malaysia.
Journal of Tropica! Ecology.< 6: 421 -431.

Frey, J.K. 2006. Inferring species distribution in the absence of
occurrence records: An example considering wolverine ( Gulo
gulo) and Canada lynx (Lynx canadensis) in New Mexico.
Biological Conservation 130: 16-24.

Hall, L.S., Richards, G.C. and Abdullah, M.T. 2002. The bats of
Niah National Park, Sarawak. Sarawak Museum Journal 78:
255-282.

Hazebroek, H.P. and Abang Morshidi, A.K. 2000. National parks
of Sarawak. Natural History Publication (Borneo): Kota
Kinabalu.

Kingston,T., Francis, C.M., Zubaid, A. and Kunz,T.H. 2003. Species
richness in an insectivorous bat assemblage from Malaysia.
Journal of Tropical Ecology 19: 67-79.

Lane, D.J.W., Kingston, T. and Lee, B.P. 2006. Dramatic decline in bat
species richness in Singapore, with implications for Southeast
Asia. Biological Conservation 131: 584-593.

Mohd-Azlan, 1, Lisa, D.P.A., Lading, E. and Mohidin, R. 2007.
Camera trapping and conservation in Kubah National Park.
Pp. 87-101. In: Unknown (ed.) Proceedings of the eighth
hornbill workshop on protected areas and biodiversity
conservation. Kuching.

Mohd-Azlan. J., Maryonto, I., Kartono, A.P. and Abdullah, M.T. 2003.
Diversity, relative abundance and conservation of chiropterans
in KayanMcnterang National Park, East Kalimantan, Indonesia.
Sarawak Museum Journal 58: 251-265.

Payne, J., Francis, C.M. and Phillips, K. 2005. A field guide to the
mammals of Borneo. The Sabah Society and WWF Malaysia:
Kota Kinabalu.

Salleh. M.A., Slim, E.U.H., Rahman, M.A. and Abdullah, M.T. 1999.
Isolation of genomic DNA from fruit bats of Kelabit Highlands
for DNA archiving and determination of genetic variation.
Pp. 179-187 In: Ismail, G. and Ali. L. (eds) A scientificjourney
through Borneo: Crocker Range National Park Sabah. ASEAN
Academic Press: Malaysia.

Struebig, M.J., Galdikas B.M.F. and Suatma. 2006. Bat diversity in
oligotrophic forests of southern Borneo. Orv.x 40: 447M55.
Tuen, A.A., Lakint. M. and Hall, L.S. 2002. Preliminary survey of
bats of the Crocker Range National Park Sabah, Malaysia.
Pp. 231-240 In: Ismail,G. and Din, L. (eds) A scientific journey
through Borneo: Bario The Kelabit Highlands of Sarawak.
Pelanduk Publications: Malaysia

Tan, K.H., Zubaid, A. and Kunz. T.H. 1998. Food habits of Cynopterus
brachyotis (Muller) (Chiroptera: Pteropodidae) in peninsular
Malaysia. Journal of Tropical Ecology> 14: 299-307.

Zar, J.H. 1996. Biostatistical analysis. Prentice Hall: New Jersey.

Accepted 27 May 2008

155

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 157

Short Communication

A previously unpublished record of Liphyra brassolis Westwood, 1864
(Lepidoptera: Lycaenidae: Miletinae) from Vella Lavella, New Georgia group,

Solomon Islands

W. JOHN TENNENT

Department of Entomology’, The Natural History’ Museum, London SW7 5BD, ENGLAND
jtstorment@googlemail.com

In transcribing letters written between 1894 and 1931 by
English explorer/naturalist/collector Albert Stewart Meek
(1871-1943) to the stafTof Rothschild’s museum at Tring,
Hertfordshire, England, references were noted regarding a
sighting (but not capture) of the miletine lycaenid butterfly
Liphyra brassolis Westwood, 1864, on the island of Vella
Lavella (New Georgia group), Solomon Islands. Liphyra
brassolis has not previously been recorded from this island
(Tennent 2002).

Liphyra brassolis occurs from northern India and
Thailand, through the Malay Peninsula, Indonesia, the
Philippines to Australia, New Guinea and the Solomon
Islands. Two subspecies are known to occur in the Solomons
Archipelago: L. brassolis bougainvilleana Samson and
Smart, 1980 (Bougainville, Papua New Guinea; Faisi,
Shortland group) and L. brassolis salomonis Samson and
Smart, 1980 (Rendova, New Georgia group; Russell group;
Guadalcanal, Florida, San Cristobal) (Tennent 2006). The
few specimens known from the Solomon Islands, and its
widespread but local distribution, almost certainly reflect the
fact that adults are seldom observed, probably due at least
in part to their crepuscular habits. Liphyra brassolis is more
likely to be found by examining the nests of the green tree
ant Oecophylla smaragdina Fabricius for larvae/pupae.

The only published record of this species from islands
of the New Georgia group is a solitary specimen caught
by A.S. Meek on Rendova in February 1904. However,
in a letter written to Rothschild’s Lepidoptera curator,
Karl Jordan, in March 1908, Meek said: “I saw one vety
fine butterfly on Vella Lavella. 1 forget the name of the
Queensland form. It is like a vety large skipper. Mr Dodd
ofKurunda published an article on its life history, in which
he says it feeds in the nests of the green tree ant. The insect
I saw was larger than either Queensland or New Guinea
forms. I think I 've taken about two specimens altogether in
latter place. It also appeared to have lighter red markings.
It was hovering over one particularly thorny tree. I 'm sony
now I didn't shoot it” (Meek 1908a).

The question of shooting the specimen, rather than the
rather more conventional use of a net, is not as bizarre as
it might sound. Several historically important birdwing
butterflies were collected by shooting them (Ackery
1997; Tennent 1997, 1999) with mustard seed or dust shot

cartridges designed for shooting small birds at short range
without causing damage to the plumage (Mearns and
Meams 1998).

Even if the description provided by Meek did not
obviously refer to Liphyra - which it clearly does -
reference to Dodd’s publications five years earlier (Dodd
1902, 1903) removes any doubt. In another letter, to Tring’s
ornithological curator Ernst Hartert later in 1908, Meek
voiced his disappointment at the meagre number of birds
collected on Vella Lavella, and added: “If I’d been able to get
that butterfly I saw (Liphyra Brassiolis [sic] of Queensland)
I should have felt more satisfied ’ (Meek 1908b).

REFERENCES

Ackery, P.R. 1997. The Natural History Museum collection of
Omithoptera (birdvving) butterflies (Lepidoptera: Papilionidae).
The Biology Curator 8: 11-17.

Dodd, F.P. 1902. Contribution to the life-history of Liphyra brassolis,
Westw., The Entomologist 35 (469): 153-156,(470): 184-188,
pi. 4.

Dodd, F.P. 1903. The young larva of Liphyra brassolis, Westw., The
Entomologist 36 (483): 211-212.

Meek, A.S. 1908a. Letter to Karl Jordan dated 29th March 1908 from
Gizo, Solomon Islands. Meek letter 207, BMNH Archives.
Unpublished.

Meek, A.S., 1908b. Letter to Ernst Hartert dated 12th August 1908
from on board the vessel “ Shamrock ”, Samarai, British New
Guinea. Meek letter 213, BMNH Archives. Unpublished.
Samson, C. and Smart, P. 1980. A review of the genus Liphyra
(Lepidoptera: Lycaenidae) of Indo-Australia, with descriptions
of two new subspecies from the Solomon Archipelago.
Aurelian, Berkley 1 (4): 6-16.

Tennent. W.J. 1997. The type locality of Omithoptera victoriae Gray,
1856, and the circumstances of the capture of the holotype
female (Lepidoptera, Ritopalocera). Archives of Natural
History 24 (2): 163-173.

Tennent, W. J. 1999. Charles Morris Woodford C.M.G. (1852-1927):
Pacific adventurer and forgotten Solomon Islands naturalist.
Archives of Natural History’ 26 (3): 419-432.

Tennent, W.J. 2002. Butterflies of the Solomon Islands: systematics
and biogeography. Storm Entomological Publications:
Dereham, England,

Tennent, W.J. 2006. A checklist of the butterflies of Melanesia,
Micronesia. Polynesia and some adjacent areas. Zootaxa
1178: 1-209

Accepted 15 October 2008

The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2008 24: 159-162

Corrigendum

Corrigendum to Horner, P. and Adams, M. (2007). A molecular systematic
assessment of species boundaries in Australian Cryptoblepharus (Reptilia:
Squamata: Scincidae) - a case study for the combined use of allozymes and
morphology to explore cryptic biodiversity. The Beagle, Records of the Museums
and Art Galleries of the Northern Territory, Supplement 3: 1-19.

PAUL HORNER

Museum and Art Gallery Northern Territory
GPO Box 4646, Darwin, NT0801, AUSTRALIA
paul.horner@nt.gov.au

In the published version of this article three tables, referred to in this text as Tables 1, 2 and 3 were omitted in error.
This error does not change the conclusions of the study in any way and the authors wish to apologise for the omission.
The paper should have contained the three following tables.

P. Homer

Table 1. Allozyme frequencies for the 11 diagnosable OTUs identified within lineage 1, plus the two extralimital species. For polymorphic
loci, the frequencies of all but the rarer/rarest alleles are expressed as percentages and shown as superscripts (allowing the frequency of each
rare allele to be calculated by subtraction from 100%). Alleles not separated by a comma all shared the frequency indicated. The following
loci were invariant: Gapd", Lap", Pganf, and Sod h .

Locus

earn

mega

plag

eger

novo

Acon-1

b w ,a

e 50 , c 25 ,b

Acon-2

c 77 ,d l5 ,b

c^.d 15 ,

c 93 ,d

c* 3 ,d

c 73 .g

c-Ad 38 . P.
b 3 ,g

c M .b 28 ,

d s ,a 2 ,g

Acp

c 97 ,e

c 97 ,a 3

c w ,f

c 97 .F,b

Acyc

c 97 ,e

c 98 ,bd'

c 98 ,e

Ada

o 88 ,n 7 ,

P.p

q 67 ,o 33

i 5i ,c",f

c 9l ,P,

P,F

o S3 ,n 21 ,

l",m 3 ,

kip 2 ,

hgrs 1

o“,

r 32 ,el'

Adh-1

b 97 ,a 2 ,c

Adh-2

e 88 ,bc\f

e 95 ,f

e 97 ,h

e 93 ,h

e 92 ,c J ,F,agl

e^.g-'.a

Alb

c“,d

c^.ab'

c",b

b 50 ,c

b 97 ,a

Dia

h 98 ,j

b“,d

d 8s ,g

b 94 ,d

P°,d",b

d 83 j

d 95 ,g\h

h 95 j

c 8l ,h 18 ,k

Enol

b 97 ,a

b w .c

b",c

Fdp

b 88 ,d 12

b 83 ,d 17

b",a

Fum

h“j

h 9l ,d

h 72 ,d

h s2 .d 13 ,

j : ,ail'

h 97 j 2 .f

h 75 ,d

Gda

e 85 ,g l3 ,h

c 70 ,e

c 88 ,e 12

e 8 \g

e 97 ,c

e %. g

c 95 ,a

Gdh

Glo

b» d »

d^c

d".a

Got-1

b 95 ,d

b 97 ,a

b' M ,d 3 ,

c 2 ^

Got-2

i 95 ,k 3 .e

m 95 ,l

g 75 ,f

P’.g

P 7 ,g

PMv’.gi 2

i 98 .ek'

i",k

Gpd

c 98 ,b

c 98 ,ab'

c",b

c 75 ,d

Gpi

a 93 ,c*,f

f»,a

P 5 ,i

P',a

a 98 ,fg'

d 9, j 2 ,i

Gpx

e 98 ,a

e 9S ,c

e r \c

e^.cd 1

Guk

c 97 ,f

a 67 ,b

F 7 ,e

P,e

c 75 ,P 2 ,e 8 .

d’.gh 1

b 82 ,d",

Hbdh

g 32 ,k 29 ,

j 7 ,m 7 ,

c\h

h«j

j 95 ,i

gM iC 25j

g^J 28 ,

be 3

d 33 ,g 33 ,

c l7 0

b-'MA'.g

c 45 ,g 28 ,

a I5 ,b l0 ,h

g 7l ,m 8 ,

h 7 .j 6 ,ko : ,

cefi'

l 9l jMi

Idh

c 97 ,a : ,d

c 50 /

P 7 ,c

P\c

p*,i

c 9J ,d 4 ,eP

C S6 ,P 2 ,be'

Ldh

Mdh-1

b 97 ,a

Mdh-2

a",c

Mpi

c 98 ,f

c*h 33 ,f

c“,f

(^“.bc 1

C 78 ,P 7 ,

d 3 ,eg'

b",a

PepA-1

e 98 ,f

e“,h

e",d

e 87 ,g

PepA-2

d 97 ,e 6 ,c

d’V

e^d 29 ,

d 93 ,c 6 ,e

e 50 ,g

PepB

d 98 ,a 2

g“i 45

b w ,d»

d 9l ,b 3 ,

g 3 .h 3

d 83,gl7

d 85 ,e 13 ,

g 2

d 87 .g 7 ,

eh 2 ,ab'

g 9 ’,h 4 .d

PepD-l

b 8 “,a 15 ,

d’.e

b 98 ,d

b % ,d\

d 67 ,e 32 ,a

a w ,d

PepD-2

d 95 ,g\b

b^.e

d“b

e^h 38 ,

b 67 ,d 17 ,

d 67 ,b 17 ,

d 92 ,g 5 ,

d s7 ,b 7 ,a 3 ,

egh'

d % ,a

6Pgd

F’.e 7 ,

hi-'.bk 2

P 5 ,h

d»,g

P',d 6 ,i

f*,i

P.bd 2 ^!'

d ?s ,f

Pgk

b 98 ,c

c^.f

e’V

el 1

c 97 ,b

b 93 .c 6 ,d

Pgm-1

c= 8 ,e 37 ,

h\f

d 70 ,c

a'V’.f

c 75 ,f

c 50 /

c 67 /

e 75 ,c

c 98 ,f

c 67 ,e 28 ,

P.a

e 93 ,d 6 ,h

Pgm-2

c 65 ^ 30 ,

a 3 .d

c*,b

b 83 ,c

b 35 ,a

b 75 ,^ 4 ^

b",e

Srdh

d 83 ,c

d 97 ,g

d 98 ,e

d",e

d 95 .g

Tpi

b 83 ,e

b 95 ,a

160

Corrigendum to Homer and Adams (2007)

Table 2. Allozyme frequencies for the 16 diagnosable OTUs identified within lineage 2, plus the putative ‘virgA 1x3 ’ hybrid population. Format
as per Table 1. The following loci were invariant: Acp-l c , AIb‘, Gapd“, Gdh“, Lap", Ldh a , Mdh-l b , Mdh-2 a , Pganf, PglP, and Sod“.

Locus

earn

fulin

horn

litor

mega

virg

''irg

A 1x3'

virg

Acon-1

d 98 ,e

d 83 ,a

d 77 .b

b 94 ,d

Acon-2

d 94 ,cP

d 9, ,f

d»c

d 9S ,f

d 92 ,c

c 70 ,d

c 55 ,d

d' w ,g 7 ,

d'",c 6 .

d s V,

be 6

d S6 ,cP,

d*b 14 ,

Acyc

c 90 ^

c 95 ,d

c 94 ,b

b^c

Ada

o M ,p 7 ,

s 4 ,ri 2 ,

l 59 ,i

1 92 ,0

j 75 ,0

o“o

o 55 j

o«F,

p’M 8 ,

js 4 ,hr',q

o w ,s

o M ,i 32 ,

0 s7 ,i

o 46 ,! 29 ,

i l5 J

Adh-1

b 98 ,d

b 98 ,a

Adh-2

P 8 ,d 6 ,

F’j

P°,e

P 5 .k 3 ,i

F 7 ,i

P‘,e

P.ci 9

P 6 ,i

F 7 ,i

c 98 ,b

b 85 .a

Dia

d !0 ,h

e 81 ,h

Enol

b 9l ,d

b 9i .d

Fdp

b”,d

b 9i .d

Fum

e 97 .b

e^.b

e 89 .g

e 97 ,g

e 85 ,c

e 97 ,c

e % ,c

e 94 ,j

c 55 ,e

e M ,c

Gda

e^h

e^g 6 ,

be 2

c 96 .e

d 88 ,f

e 92 ,d 9 ,f

e w ,f

Glo

d^.a

d 7S ,a

d 98 ,a

a H .d

d 8l ',ae 7

Got-1

b' M .c

b 50 ,c

b ,5 .d

b 67 ,c l7 ,d

b*,ad ;

b 94 .a

b 93 .d

Got-2

d w .h

g“,h“i

g 98 ,i

d 94 ,b

i 86 .g

g 7l .i”,d

g*j

Gpd

c 96 ,b

Gpi

a*,tlf

P,a

a 91 ,e 6 ,h

a % .bg :

a 9l ,d 5 ,g

Gpx

e^.b

b“,e

e s) ,f

e 93 .b

Guk

e^h

h w .e

Hbdh

tF.d

m 8 “,h

h M ,d

h 90 .!

j*h 3J ,

k i5 ,p 4 ,q

i*,h

p 75 ,m

m*,h

m^.dh 3

m 52 ,o 46 ,

o 8l .m l4 ,p

j so ,h 3! ,

o 9 ,k'.m

o 50 j 79 ,m

m 69 ,o

Idh

c 9 \h

c 98 ,a

Mpi

c 9! ,f

c 80 ,b

c 98 ,f

c*',P 6 ,b

c 93 .f

c 92 ,f

PepA-1

b 9l ,e

c w ,a

e 50 ,b 40 ,d

e 98 ,d

e"',g

e 50 /

e 66 ^

e*b

e 87 ,b

e 9l ,b

e 79 ,b

e 96 .d

PepA-2

e“,f

e 93 ,c 5 ,d

e 80 ,d 10 ,c

d 5 “,e

e»f

e 97 .c

b 8l .c IJ ,

a“,d

b 7S ,a

e 55 /-,

d l8 ,c

dV 1 ,

e l4 ,b

PepB

g 90 ,!

g 70 ,c

g”.h 15 ,

dfi 2

g ,J .d

g 75 ,b

g 80 ,d

d 85 ,g

h S8 ,dfjk 5

g 7l ,h :7 ,f

g 7S .h

g“,h :9 .d

g 96 ,h

PepD-l

d 97 ,e

dT,h

d 96 ,be 2

d 79 .g

d 84 ,g

d 98 ,e

c 44 .g 5l ,d

d 95 ,e

d 93 ,e

PepD-2

d 79 ,g

d^e

d' M ,i

d 97 .a

d 69 ,c

d 95 .g

6Pgd

P 7 ,d 20 ,i

F°,bdg 10

f«i

P‘',d 37 ,

b 7 ,gj J ,i

P,i

f“.i 7 j

d 94 ,aci !

P 7 ,d

d 70 ,f

d^f

Pgm-1

d 79 ,g

d«,a 41 ,

g 9 ,e

d*g

c^.d

cV°,g

d 67 ,c

d 9! ,b

d»,g

d !l ,c l3 ,g

d 8 ",g l5 ,a

d 79 ,a l4 ,g

d 92 ,g

Pgm-2

c 98 ,d

c 97 ,b

c 96 ,e

c 88 ,e

c 87 .d

c 95 .d

c 86 ,d

Srdh

c 97 ,f

c 70 ,a

c 96 ,b

c 9J ,a

c 90 ^

c 77 ,b

Tpi

a 7: ,b :s ,d

a 96 ,b

a 93 ,c

161

P. Homer

Table 3. Pairwise genetic distance measures among all final OTUs. OTUs are grouped and delineated according their membership of either
lineage 1 (top), lineage 2 (middle) or extralimital taxa (bottom). Lower triangle = %FD; upper triangle = Nei D.

162

*3,-11S2.

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Brake, B., McNeish, J. and Simmons, D. 1979. Art of the Pacific. Oxford University Press: Wellington, New Zealand.

Callcn, R.A. 1984. Clays of the palygorskite-sepiolitc group: depositional environment, age and distribution. Pp 1-38. In: Singer, A. and
Galan, E. (eds) Palygorskite-sepiolite occurrence, genesis ami uses. Elsevier: Amsterdam.

Crowley, L.M. 1949. Working class conditions in Australia, 1788-1851. Unpublished PhD thesis. University of Melbourne.

Sadlier, R.A. 1990. A new species of scincid lizard from western Arnhem Land, Northern Territory. The Beagle, Records of the Northern
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The Beagle

Records of the Museums and Art Galleries
of the Northern Territory

Volume 24, December 2008
CONTENTS

BERRA, T. M. - Charles Darwin’s paradigm shift.1

DURETTO, M. F. - A reassessment of Boronia (Rutaceae) in the Northern Territory with a key to species,

the description of one new species and the reduction, in synonymy, of another species.7

PAXTON, H. and SAFARIK, M. - Jaw growth and replacement in Diopatra aciculata (Annelida: Onuphidae).15

SALGADO KENT, C. P. and McGUINNESS, K. A. - Feeding selectivity of sesarmid crabs from

northern Australian mangrove forests.’.23

BRABY, M. F. - Taxonomic review of Candalides absimilis (C. Felder, 1862) and C. margarita (Semper, 1879)
(Lepidoptera: Lycaenidae), with descriptions of two new subspecies.33

BAEHR, M. - Two new species of the genus Pseudaptinus Castelnau from northern Australia

(Insecta: Coleoptera: Carabidae: Zuphiinae).55

KOTT, P. - Biogeographic implications of Ascidiacea (Tunicata) from the Wessel Islands (Arafura Sea).63

THORBURN, D. C. and ROWLAND, A. J. — Juvenile bull sharks Carcharhinus leucas (Valenciennes, 1839)

in northern Australian rivers. 79

M0LLER, P. R. and SCHWARZHANS, W. - Review of the Dinematichthyni (Teleostei: Bythitidae)

of the Indo-west Pacific. Part IV. Dinematichthys and two new genera with descriptions of nine new species.87

LARSON, H. K. - Anew species of the gudgeon Bostiychus (Teleostei: Gobioidei: Eleotridae), from

peninsular Malaysia.147

MOHD-AZLAN, J., TAHA, S. H., LAMAN, C. J. M. and ABDULLAH, M. T. - Diversity of bats at

two contrasting elevations in a protected dipterocarp forest in Sarawak, Borneo.151

Short communication

TENNENT, W. J. - A previously unpublished record of Liphyra brassolis Westwood, 1864

(Lepidoptera: Lycaenidae: Miletinae) from Vella Lavella, New Georgia group, Solomon Islands.157

Corrigendum

HORNER, P. - Corrigendum to Horner, P. and Adams, M. (2007). A molecular systematic assessment of
species boundaries in Australian Cryptoblepharus (Reptilia: Squamata: Scincidae) - a case study for the
combined use of allozymes and morphology to explore cryptic biodiversity. The Beagle, Records of the
Museums and Art Galleries of the Northern Territory, Supplement 3: 1-19.159

ABSTRACTED IN ZOOLOGICAL RECORD AND BIOSIS

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